Latitudinal gradients in species diversity

Species richness, or biodiversity, increases from the poles to the tropics for a wide variety of terrestrial and marine organisms, often referred to as the latitudinal diversity gradient (LDG)[1]. The LDG is one of the most widely recognized patterns in ecology[1]. The LDG has been observed to varying degrees in Earth's past.[2] A parallel trend has been found with elevation (elevational diversity gradient)[3], though this is less well-studied[4]

Explaining the latitudinal diversity gradient has been called one of the great contemporary challenges of biogeography and macroecology (Willig et al. 2003, Pimm and Brown 2004, Cardillo et al. 2005).[5] The question "What determines patterns of species diversity?" was among the 25 key research themes for the future identified in 125th Anniversary issue of Science (July 2005). There is a lack of consensus among ecologists about the mechanisms underlying the pattern, and many hypotheses have been proposed and debated. A recent review [6] noted that among the many conundrums associated with the LDG (or LBG, Latitudinal Biodiversity Gradient) the causal relationship between rates of molecular evolution and speciation has yet to be demonstrated.

Understanding the global distribution of biodiversity is one of the most significant objectives for ecologists and biogeographers. Beyond purely scientific goals and satisfying curiosity, this understanding is essential for applied issues of major concern to humankind, such as the spread of invasive species, the control of diseases and their vectors, and the likely effects of global climate change on the maintenance of biodiversity (Gaston 2000). Tropical areas play prominent roles in the understanding of the distribution of biodiversity, as their rates of habitat degradation and biodiversity loss are exceptionally high.[7]

Patterns in the past

The LDG is a noticeable pattern among modern organisms that has been described qualitatively and quantitatively. It has been studied at various taxonomic levels, through different time periods and across many geographic regions (Crame 2001). The LDG has been observed to varying degrees in Earth's past, possibly due to differences in climate during various phases of Earth's history. Some studies indicate that the LDG was strong, particularly among marine taxa, while other studies of terrestrial taxa indicate the LDG had little effect on the distribution of animals.[2]

Hypotheses for pattern

Although many of the hypotheses exploring the latitudinal diversity gradient are closely related and interdependent, most of the major hypotheses can be split into three general hypotheses.

Spatial/Area hypotheses

There are five major hypotheses that depend solely on the spatial and areal characteristics of the tropics.

Mid-domain effect

Using computer simulations, Colwell and Hurtt (1994) and Willig and Lyons (1998) first pointed out that if species’ latitudinal ranges were randomly shuffled within the geometric constraints of a bounded biogeographical domain (e.g. the continents of the New World, for terrestrial species), species' ranges would tend to overlap more toward the center of the domain than towards its limits, forcing a mid-domain peak in species richness. Colwell and Lees (2000) called this stochastic phenomenon the mid-domain effect (MDE), presented several alternative analytical formulations for one-dimensional MDE (expanded by Connolly 2005), and suggested the hypothesis that MDE might contribute to the latitudinal gradient in species richness, together with other explanatory factors considered here, including climatic and historical ones. Because "pure" mid-domain models attempt to exclude any direct environmental or evolutionary influences on species richness, they have been claimed to be null models (Colwell et al. 2004, 2005). On this view, if latitudinal gradients of species richness were determined solely by MDE, observed richness patterns at the biogeographic level would not be distinguishable from patterns produced by random placement of observed ranges (Colwell and Lees 2000). Others object that MDE models so far fail to exclude the role of environment at the population level and in setting domain boundaries, and therefore cannot be considered null models (Hawkins and Diniz-Filho 2002; Hawkins et al. 2005; Zapata et al. 2003, 2005). Mid-domain effects have proven controversial (e.g. Jetz and Rahbek 2001, Koleff and Gaston 2001, Lees and Colwell, 2007, Romdal et al. 2005, Rahbek et al. 2007, Storch et al. 2006; Bokma and Monkkonen 2001, Diniz-Filho et al. 2002, Hawkins and Diniz-Filho 2002, Kerr et al. 2006, Currie and Kerr, 2007). While some studies have found evidence of a potential role for MDE in latitudinal gradients of species richness, particularly for wide-ranging species (e.g. Jetz and Rahbek 2001, Koleff and Gaston 2001, Lees and Colwell, 2007, Romdal et al. 2005, Rahbek et al. 2007, Storch et al. 2006; Dunn et al. 2007)[5][8] others report little correspondence between predicted and observed latitudinal diversity patterns (Bokma and Monkkonen 2001, Currie and Kerr, 2007, Diniz-Filho et al. 2002, Hawkins and Diniz-Filho 2002, Kerr et al. 2006).

Geographical area hypothesis

Another spatial hypothesis is the geographical area hypothesis (Terborgh 1973). It asserts that the tropics are the largest biome and that large tropical areas can support more species. More area in the tropics allows species to have larger ranges, and consequently larger population sizes. Thus, species with larger ranges are likely to have lower extinction rates (Rosenzweig 2003). Additionally, species with larger ranges may be more likely to undergo allopatric speciation, which would increase rates of speciation (Rosenzweig 2003). The combination of lower extinction rates and high rates of speciation leads to the high levels of species richness in the tropics.

A critique of the geographical area hypothesis is that even if the tropics is the most extensive of the biomes, successive biomes north of the tropics all have about the same area. Thus, if the geographical area hypothesis is correct these regions should all have approximately the same species richness, which is not true, as is referenced by the fact that polar regions contain fewer species than temperate regions (Gaston and Blackburn 2000). To explain this, Rosenzweig (1992) suggested that if species with partly tropical distributions were excluded, the richness gradient north of the tropics should disappear. Blackburn and Gaston 1997 tested the effect of removing tropical species on latitudinal patterns in avian species richness in the New World and found there is indeed a relationship between the land area and the species richness of a biome once predominantly tropical species are excluded. Perhaps a more serious flaw in this hypothesis is some biogeographers suggest that the terrestrial tropics are not, in fact, the largest biome, and thus this hypothesis is not a valid explanation for the latitudinal species diversity gradient (Rohde 1997, Hawkins and Porter 2001). In any event, it would be difficult to defend the tropics as a "biome" rather than the geographically diverse and disjunct regions that they truly include.

The effect of area on biodiversity patterns has been shown to be scale dependent, having the strongest effect among species with small geographical ranges compared to those species with large ranges who are affected more so by other factors such as the mid-domain and/or temperature.[5]

Species-energy hypothesis

The species energy hypothesis suggests the amount of available energy sets limits to the richness of the system. Thus, increased solar energy (with an abundance of water) at low latitudes causes increased net primary productivity (or photosynthesis). This hypothesis proposes the higher the net primary productivity the more individuals can be supported, and the more species there will be in an area. Put another way, this hypothesis suggests that extinction rates are reduced towards the equator as a result of the higher populations sustainable by the greater amount of available energy in the tropics. Lower extinction rates lead to more species in the tropics.

One critique of this hypothesis has been that increased species richness over broad spatial scales is not necessarily linked to increased number of individuals, which in turn is not necessarily related to increased productivity.[9] Additionally, the observed changes in the number of individuals in an area with latitude or productivity are either too small (or in the wrong direction) to account for the observed changes in species richness.[9] The potential mechanisms underlying the species-energy hypothesis, their unique predictions and empirical support have been assessed in a major review by Currie et al. (2004).[10]

The effect of energy has been supported by several studies in terrestrial and marine taxa [7]

Climate harshness hypothesis

Another climate-related hypothesis is the climate harshness hypothesis, which states the latitudinal diversity gradient may exist simply because fewer species can physiologically tolerate conditions at higher latitudes than at low latitudes because higher latitudes are often colder and drier than tropical latitudes. Currie et al. (2004)[10] found fault with this hypothesis by stating that, although it is clear that climatic tolerance can limit species distributions, it appears that species are often absent from areas whose climate they can tolerate.

Climate stability hypothesis

Similarly to the climate harshness hypothesis, climate stability is suggested to be the reason for the latitudinal diversity gradient. The mechanism for this hypothesis is that while a fluctuating environment may increase the extinction rate or preclude specialization, a constant environment can allow species to specialize on predictable resources, allowing them to have narrower niches and facilitating speciation. The fact that temperate regions are more variable both seasonally and over geological timescales (discussed in more detail below) suggests that temperate regions are thus expected to have less species diversity than the tropics.

Critiques for this hypothesis include the fact that there are many exceptions to the assumption that climate stability means higher species diversity. For example, low species diversity is known to occur often in stable environments such as tropical mountaintops. Additionally, many habitats with high species diversity do experience seasonal climates, including many tropical regions that have highly seasonal rainfall (Brown and Lomolino 1998).

Historical/Evolutionary hypotheses

There are three main hypotheses that are related to historical and evolutionary explanations for the increase of species diversity towards the equator.

The historical perturbation hypothesis

The historical perturbation hypothesis proposes the low species richness of higher latitudes is a consequence of an insufficient time period available for species to colonize or recolonize areas because of historical perturbations such as glaciation (Brown and Lomolino 1998, Gaston and Blackburn 2000). This hypothesis suggests that diversity in the temperate regions have not yet reached equilibrium, and that the number of species in temperate areas will continue to increase until saturated (Clarke and Crame 2003).

The evolutionary rate hypothesis

The evolutionary rate hypothesis argues higher evolutionary rates in the tropics have caused higher speciation rates and thus increased diversity at low latitudes (Cardillo et al. 2005, Weir & Schluter 2007, Rolland et al. 2014). Higher evolutionary rates in the tropics have been attributed to higher ambient temperatures, higher mutation rates, shorter generation time and/or faster physiological processes (Rohde 1992, Allen et al. 2006), and increased selection pressure from other species that are themselves evolving[11]. Faster rates of microevolution in warm climates (i.e. low latitudes and altitudes) have been shown for plants (Wright et al. 2006), mammals (Gillman et al. 2009) and amphibians (Wright et al. 2010). Based on the expectation that faster rates of microevolution result in faster rates of speciation, these results suggest that faster evolutionary rates in warm climates almost certainly have a strong influence on the latitudinal diversity gradient. More research needs to be done to determine whether or not speciation rates actually are higher in the tropics. Understanding whether extinction rate varies with latitude will also be important to whether or not this hypothesis is supported (Rolland et al. 2014).

The hypothesis of effective evolutionary time

The hypothesis of effective evolutionary time assumes that diversity is determined by the evolutionary time under which ecosystems have existed under relatively unchanged conditions, and by evolutionary speed directly determined by effects of environmental energy (temperature) on mutation rates, generation times, and speed of selection (Rohde 1992). It differs from most other hypotheses in not postulating an upper limit to species richness set by various abiotic and biotic factors, i.e., it is a nonequilibrium hypothesis assuming a largely non-saturated niche space. It does accept that many other factors may play a role in causing latitudinal gradients in species richness as well. The hypothesis is supported by much recent evidence, in particular the studies of Allen et al. (2006) and Wright et al. (2006).

Biotic hypotheses

Biotic hypotheses claim ecological species interactions such as competition, predation, mutualism, and parasitism are stronger in the tropics and these interactions promote species coexistence and specialization of species, leading to greater speciation in the tropics. These hypotheses are problematic because they cannot be the proximate cause of the latitudinal diversity gradient as they fail to explain why species interactions might be stronger in the tropics. An example of one such hypothesis is the greater intensity of predation and more specialized predators in the tropics has contributed to the increase of diversity in the tropics (Pianka 1966). This intense predation could reduce the importance of competition (see competitive exclusion) and permit greater niche overlap and promote higher richness of prey. While recent large-scale experiments suggest predation may be more intense in the tropics[12][13], this cannot be the ultimate cause of high tropical diversity because it fails to explain what gives rise to the richness of the predators in the tropics.

Several recent studies have failed to observe consistent changes in ecological interactions with latitude (Lambers et al. 2002)[1]. These studies suggest the intensity of species interactions are not correlated with the change in species richness with latitude.

Synthesis and conclusions

There are many other hypotheses related to the latitudinal diversity gradient, but the above hypotheses are a good overview of the major ones still cited today. It is important to note that many of these hypotheses are similar to and dependent on one another. For example, the evolutionary hypotheses are closely dependent on the historical climate characteristics of the tropics.

The generality of the latitudinal diversity gradient

An extensive meta-analysis of nearly 600 latitudinal gradients from published literature tested the generality of the latitudinal diversity gradient across different organismal, habitat and regional characteristics[1]. The results showed that the latitudinal gradient occurs in marine, terrestrial, and freshwater ecosystems, in both hemispheres. The gradient is steeper and more pronounced in richer taxa (i.e. taxa with more species), larger organisms, in marine and terrestrial versus freshwater ecosystems, and at regional versus local scales. The gradient steepness (the amount of change in species richness with latitude) is not influenced by dispersal, animal physiology (homeothermic or ectothermic) trophic level, hemisphere, or the latitudinal range of study. The study could not directly falsify or support any of the above hypotheses, however results do suggest a combination of energy/climate and area processes likely contribute to the latitudinal species gradient. Notable exceptions to the trend include the ichneumonidae, shorebirds, penguins, and freshwater zooplankton.

Data robustness

One of the main assumptions about LDGs and patterns in species richness is that the underlying data (i.e. the lists of species at specific locations) are complete. However, this assumption is not met in most cases. For instance, diversity patterns for blood parasites of birds suggest higher diversity in tropical regions, however, the data may be skewed by undersampling in rich faunal areas such as Southeast Asia and South America.[14] For marine fishes, which are among the most studied taxonomic groups, current lists of species are considerably incomplete for most of the world's oceans. At a 3° (about 350 km2) spatial resolution, less than 1.8% of the world's oceans have above 80% of their fish fauna currently described.[15]

Conclusion

The fundamental macroecological question that the latitudinal diversity gradient depends on is ‘What causes patterns in species richness?'. Species richness ultimately depends on whatever proximate factors are found to affect processes of speciation, extinction, immigration, and emigration. While some ecologists continue to search for the ultimate primary mechanism that causes the latitudinal richness gradient, many ecologists suggest instead this ecological pattern is likely to be generated by several contributory mechanisms (Gaston and Blackburn 2000, Willig et al. 2003, Rahbek et al. 2007). For now the debate over the cause of the latitudinal diversity gradient will continue until a groundbreaking study provides conclusive evidence or there is general consensus that multiple factors contribute to the pattern.

See also

References

  1. ^ a b c d Hillebrand, Helmut (2004). "On the Generality of the Latitudinal Diversity Gradient". The American Naturalist. 163 (2): 192–211. doi:10.1086/381004. ISSN 0003-0147. PMID 14970922.
  2. ^ a b Sahney, S.; Benton, M.J. (2008). "Recovery from the most profound mass extinction of all time" (PDF). Proceedings of the Royal Society B. 275 (1636): 759–65. doi:10.1098/rspb.2007.1370. PMC 2596898. PMID 18198148.
  3. ^ McCain, Christy M. (2005). "ELEVATIONAL GRADIENTS IN DIVERSITY OF SMALL MAMMALS". Ecology. 86 (2): 366–372. doi:10.1890/03-3147. ISSN 0012-9658.
  4. ^ Rahbek, Carsten (1995). "The elevational gradient of species richness: a uniform pattern?". Ecography. 18 (2): 200–205. doi:10.1111/j.1600-0587.1995.tb00341.x. ISSN 1600-0587.
  5. ^ a b c Mora C & Robertson DR; Robertson (2005). "Causes of latitudinal gradients in species richness: a test with fishes of the Tropical Eastern Pacific" (PDF). Ecology. 86 (7): 1771–1792. doi:10.1890/04-0883.
  6. ^ Dowle, E. J.; Morgan-Richards, M.; Trewick, S. A. (2013). "Molecular evolution and the latitudinal biodiversity gradient". Heredity. 110 (6): 501–510. doi:10.1038/hdy.2013.4. PMC 3656639. PMID 23486082.
  7. ^ a b Tittensor D.; et al. (2011). "Global patterns and predictors of marine biodiversity across taxa" (PDF). Nature. 466 (7310): 1098–1101. doi:10.1038/nature09329. PMID 20668450.
  8. ^ Mora C; et al. (2003). "Patterns and processes in reef fish diversity" (PDF). Nature. 421 (6926): 933–936. doi:10.1038/nature01393. PMID 12606998.
  9. ^ a b Cardillo, M.; Orme, C. D. L.; Owens, I. P. F. (2005). "Testing for latitudinal bias in diversification rates: An example using New World birds". Ecology. 86 (9): 2278–2287. doi:10.1890/05-0112.
  10. ^ a b Currie, D. J.; Mittelbach, G. G.; Cornell, H. V.; Kaufman, D. M.; Kerr, J. T.; Oberdorff, T. (2004). "Predictions and tests of climate-based hypotheses of broad-scale variation in taxonomic richness". Ecology Letters. 7 (11): 1121–1134. doi:10.1111/j.1461-0248.2004.00671.x.
  11. ^ Schemske, Douglas W.; Mittelbach, Gary G.; Cornell, Howard V.; Sobel, James M.; Roy, Kaustuv (2009). "Is There a Latitudinal Gradient in the Importance of Biotic Interactions?". Annual Review of Ecology, Evolution, and Systematics. 40 (1): 245–269. doi:10.1146/annurev.ecolsys.39.110707.173430. ISSN 1543-592X.
  12. ^ Roslin, Tomas; Hardwick, Bess; Novotny, Vojtech; Petry, William K.; Andrew, Nigel R.; Asmus, Ashley; Barrio, Isabel C.; Basset, Yves; Boesing, Andrea Larissa (2017-05-19). "Higher predation risk for insect prey at low latitudes and elevations". Science. 356 (6339): 742–744. doi:10.1126/science.aaj1631. ISSN 0036-8075. PMID 28522532.
  13. ^ Hargreaves, A. L.; Suárez, Esteban; Mehltreter, Klaus; Myers-Smith, Isla; Vanderplank, Sula E.; Slinn, Heather L.; Vargas-Rodriguez, Yalma L.; Haeussler, Sybille; David, Santiago (2019). "Seed predation increases from the Arctic to the Equator and from high to low elevations". Science Advances. 5 (2): eaau4403. doi:10.1126/sciadv.aau4403. ISSN 2375-2548. PMC 6382403. PMID 30801010.
  14. ^ Clark, Nicholas; Clegg, S.; Lima, M. (2014). "A review of global diversity in avian haemosporidians (Plasmodium and Haemoproteus: Haemosporida): new insights from molecular data". International Journal for Parasitology. 44 (5): 329–338. doi:10.1016/j.ijpara.2014.01.004. PMID 24556563.
  15. ^ Mora C; et al. (2007). "The completeness of taxonomic inventories for describing the global diversity and distribution of marine fishes" (PDF). Proceedings of the Royal Society B. 275 (1631): 149–155. doi:10.1098/rspb.2007.1315. PMC 2596190. PMID 17999950.
Bacterivore

Bacterivores are free-living, generally heterotrophic organisms, exclusively microscopic, which obtain energy and nutrients primarily or entirely from the consumption of bacteria. Many species of amoeba are bacterivores, as well as other types of protozoans. Commonly, all species of bacteria will be prey, but spores of some species, such as Clostridium perfringens, will never be prey, because of their cellular attributes.

Copiotroph

A copiotroph is an organism found in environments rich in nutrients, particularly carbon. They are the opposite to oligotrophs, which survive in much lower carbon concentrations.

Copiotrophic organisms tend to grow in high organic substrate conditions. For example, copiotrophic organisms grow in Sewage lagoons. They grow in organic substrate conditions up to 100x higher than oligotrophs.

Decomposer

Decomposers are organisms that break down dead or decaying organisms, and in doing so, they carry out the natural process of decomposition. Like herbivores and predators, decomposers are heterotrophic, meaning that they use organic substrates to get their energy, carbon and nutrients for growth and development. While the terms decomposer and detritivore are often interchangeably used, detritivores must ingest and digest dead matter via internal processes while decomposers can directly absorb nutrients through chemical and biological processes hence breaking down matter without ingesting it. Thus, invertebrates such as earthworms, woodlice, and sea cucumbers are technically detritivores, not decomposers, since they must ingest nutrients and are unable to absorb them externally.

Dominance (ecology)

Ecological dominance is the degree to which a taxon is more numerous than its competitors in an ecological community, or makes up more of the biomass.

Most ecological communities are defined by their dominant species.

In many examples of wet woodland in western Europe, the dominant tree is alder (Alnus glutinosa).

In temperate bogs, the dominant vegetation is usually species of Sphagnum moss.

Tidal swamps in the tropics are usually dominated by species of mangrove (Rhizophoraceae)

Some sea floor communities are dominated by brittle stars.

Exposed rocky shorelines are dominated by sessile organisms such as barnacles and limpets.

Effective evolutionary time

The hypothesis of effective evolutionary time attempts to explain gradients, in particular latitudinal gradients, in species diversity. It was originally named "time hypothesis".

Feeding frenzy

In ecology, a feeding frenzy occurs when predators are overwhelmed by the amount of prey available. For example, a large school of fish can cause nearby sharks, such as the lemon shark, to enter into a feeding frenzy. This can cause the sharks to go wild, biting anything that moves, including each other or anything else within biting range. Another functional explanation for feeding frenzy is competition amongst predators. This term is most often used when referring to sharks or piranhas. It has also been used as a term within journalism.

Klaus Rohde

Klaus Rohde (born 1932 in Brandenburg an der Havel, Germany) is a German biologist at the University of New England (UNE), Australia, known particularly for his work on marine parasitology, evolutionary ecology/zoogeography, and phylogeny/ultrastructure of lower invertebrates.

Macroecology

Macroecology is the subfield of ecology that deals with the study of relationships between organisms and their environment at large spatial scales to characterise and explain statistical patterns of abundance, distribution and diversity. The term was coined by James Brown of the University of New Mexico and Brian Maurer of Michigan State University in a 1989 paper in Science.Macroecology approaches the idea of studying ecosystems using a "top down" approach. It seeks understanding through the study of the properties of the system as a whole; Kevin Gaston and Tim Blackburn make the analogy to seeing the forest for the trees.Macroecology examines how global development in climate change affect wildlife populations. Classic ecological questions amenable to study through the techniques of macroecology include questions of species richness, latitudinal gradients in species diversity, the species-area curve, range size, body size, and species abundance. For example, the relationship between abundance and range size (why species that maintain large local population sizes tend to be widely distributed, while species that are less abundant tend to have restricted ranges) has received much attention.

Mesotrophic soil

Mesotrophic soils are soils with a moderate inherent fertility. An indicator of soil fertility is its base status, which is expressed as a ratio relating the major nutrient cations (calcium, magnesium, potassium and sodium) found there to the soil's clay percentage. This is commonly expressed in hundredths of a mole of cations per kilogram of clay, i.e. cmol (+) kg−1 clay.

Mycotroph

A mycotroph is a plant that gets all or part of its carbon, water, or nutrient supply through symbiotic association with fungi. The term can refer to plants that engage in either of two distinct symbioses with fungi:

Many mycotrophs have a mutualistic association with fungi in any of several forms of mycorrhiza. The majority of plant species are mycotrophic in this sense. Examples include Burmanniaceae.

Some mycotrophs are parasitic upon fungi in an association known as myco-heterotrophy.

Organotroph

An organotroph is an organism that obtains hydrogen or electrons from organic substrates. This term is used in microbiology to classify and describe organisms based on how they obtain electrons for their respiration processes. Some organotrophs such as animals and many bacteria, are also heterotrophs. Organotrophs can be either anaerobic or aerobic.

Antonym: Lithotroph, Adjective: Organotrophic.

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.

Recruitment (biology)

In biology, especially marine biology, recruitment occurs when a juvenile organism joins a population, whether by birth or immigration, usually at a stage whereby the organisms are settled and able to be detected by an observer.There are two types of recruitment: closed and open.In the study of fisheries, recruitment is "the number of fish surviving to enter the fishery or to some life history stage such as settlement or maturity".

Relative abundance distribution

In the field of ecology, the relative abundance distribution (RAD) or species abundance distribution describes the relationship between the number of species observed in a field study as a function of their observed abundance. The graphs obtained in this manner are typically fitted to a Zipf–Mandelbrot law, the exponent of which serves as an index of biodiversity in the ecosystem under study.

Species diversity

Species diversity is the number of different species that are represented in a given community (a dataset). The effective number of species refers to the number of equally abundant species needed to obtain the same mean proportional species abundance as that observed in the dataset of interest (where all species may not be equally abundant). Species diversity consists of three components: species richness, taxonomic or phylogenetic diversity and species evenness. Species richness is a simple count of species, taxonomic or phylogenetic diversity is the genetic relationship between different groups of species,whereas species evenness quantifies how equal the abundances of the species are.

Species richness

Species richness is the number of different species represented in an ecological community, landscape or region. Species richness is simply a count of species, and it does not take into account the abundances of the species or their relative abundance distributions. Species diversity takes into account both species richness and species evenness.

Thorson's rule

Thorson's rule (named after Gunnar Thorson by S. A. Mileikovsky in 1971)

is an ecogeographical rule which states that benthic marine invertebrates at low latitudes tend to produce large numbers of eggs developing to pelagic (often planktotrophic [plankton-feeding]) and widely dispersing larvae, whereas at high latitudes such organisms tend to produce fewer and larger lecithotrophic (yolk-feeding) eggs and larger offspring, often by viviparity or ovoviviparity, which are often brooded.

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