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Latitudinal patterns


The latitudinal cline of the diversity (i.e. the highest numbers of species in the tropics and gradual decrease poleward) is a well recognized and long established pattern in terrestrial ecology. It was documented e.g. for plants, bats, mammalian quadrupeds, reptiles or termites [1]. A similar pattern has been postulated for marine biota. The first suggestion of the latitudinal diversity cline was formulated in late 1950ies for hard bottom epifauna[2]. Sanders in his influential paper[3], postulated that the macrobenthos is more diverse in the tropical shallow seas than that in boreal locations. He explained that difference by his ‘time-stability theory’ – he perceived tropics as ‘biologically accommodated’ systems that experienced stable conditions for a long time, while boreal communities as ‘physically controlled’ by environmental stressors.

Documented patterns

Clear gradients of increased diversity from the North Pole to the tropics were recorded for prosobranch mollusks collected along the Pacific and Atlantic coasts of North America[4]. Similar pattern was described for bivalves of North Pacific continental shelf[5]. A latitudinal cline in regional species richness (data pooled into bins of 10 degrees of latitude) was described for North Atlantic Bryozoa[6]. Deep-sea benthic foraminifera sample species richness decreased with increasing latitude in both North and South Atlantic[7]. Rex et al.[8] described a clear latitudinal cline for deep-sea isopods, gastropods and bivalves in North Atlantic.

Causal mechanisms

Several hypothesis have been proposed to explain the origins of the latitudinal cline of diversity in the sea[4][9][10]. It was postulated that the decrease of the diversity towards higher latitudes can be explained by:

  • time-stability hypothesis (sea Introduction)
  • Rapoport’s rule (latitudinal differences in species range sizes) - smaller geographic ranges in the tropics permit more species to co-occur in a given area, broader geographic ranges of species in higher latitudes result in lower diversities
  • species-area hypothesis – based on the species-area relationship
  • differences in geological history
  • gradients in solar energy input/levels of productivity
  • ‘mid-domain effect’ - which assumes denser species packing around the geographic midpoint of distribution
  • differences in rates of evolutionary processes (speciation and extinction)

How general is the pattern in the sea?

Some authors questioned the generality of the latitudinal pattern of diversity in the sea:

  • Thorson[2] claimed that latitudinal diversity gradient occurs in marine epifauna but not in infauna and attributed that difference to contrasting levels of heterogeneity in infaunal vs epifaunal habitats,
  • Clarke[11] noted that the pattern was documented mostly for taxa with a calcareous skeleton (Foraminifera, Bivalvia, Gastropoda) – he suggested that the low species richness in cold polar waters might be a pattern specific to calcifying taxa (resulting from higher energetic costs of calcification in low temperatures)
  • no latitudinal effect was noted for protobrach bivalves in north-eastern Pacific as opposed to clear patterns for other bivalve groups - Roy et al.[12] related these contrasts to differences in reproductive and developmental strategies employed by the studied taxa (planktotrophic vs non-planktotrophic mode of dispersal)
  • several authors documented asymmetric patterns in studies encompassing both hemispheres: a sharp cline in the northern hemisphere contrasted with no clear gradient in the southern seas was observed e.g. for deep-sea gastropods, isopods and bivalves[13] – these findings suggested that the observed latitudinal patterns could be produced by the impoverishment of north Atlantic and Arctic high latitude seas (that are considered to remain in a state of colonization after the last glaciations) rather than some globally operating processes
  • no latitudinal effect was recorded in infaunal materials encompassing: 1) three selected localities in tropical (Java), temperate (North Sea) and polar (Svalbard) waters[14], 2) Norwegian continental shelf[15], 3) European continental shelf waters spanning from 36 to 81°N – comprising Marbeff database[16].
  • no even latitudinal cline of diversity was observed for sessile macrofauna colonizing intertidal boulder fields in west North Atlantic, the diversity was only strongly depressed in High Arctic sites that was related to severe ice disturbance in littoral zonein in the Arctic[17]

Other large scale patterns of marine diversity

Longitudinal pattern

Figure 1: Distribution of coral, mangrove and seagrass diversity. (2002). In UNEP/GRID-Arendal Maps and Graphics Library. Retrieved 12:15, March 18, 2009 from [1]

A comprehensive studies of diversity of reef-building corals, bivalves and other taxa showed that the diversity of those groups is highest in the southern China-Indonesia-NE Australia region and decreases as one gets farther away (both longitudinally and attitudinally) from that area (Fig. 1). Crame[10] showed that for bivalves the steepest gradients (both longitudinal and latitudinal) were recorded for the youngest clades. That finding supports the hypothesis that the south-east Asian tropical high-diversity area served as major center of speciation and the observed gradients represented an evolutionary radiation towards other tropical regions and higher latitudes.

Depth gradients

Marine environment differs from the terrestrial systems in that it has an important third dimension – depth. Sanders[3] suggested that deep ocean provides more stable environmental conditions and so hosts a more diverse ‘biologically accommodated’ benthic biota. Grassle & Maciolek[18] reported as much as 698 species in their study of 90 000 benthic invertebrates collected off east US coast at 1500-2100 m. The parabolic pattern (with highest diversities at intermediate depths of 2000-3000 m and lower values on the upper slope and in the abyss) was documented for polychaetes, gastropods, protobranchs, cumaceans, invertebrate megafauna and fish megafauna in northwestern Atlantic and seemed to be a predominating depth related pattern in the seas[19]. That pattern was related to bathymetric changes in average population growth rates and competitive replacement (regulated by predation, disturbance level and productivity)[20]. The increase of macrobenthic diversity towards the intermediate (2000-3000 m) depths was not observed e.g. in the Greenland Sea where the ecological patterns are masked by the historical and geographical constraints[21].


  1. Rosenzweig ML (1995) Species diversity in space and time. Cambridge University Press, Cambridge
  2. 2,0 2,1 Thorson G (1957) Bottom Communities (sublittoral or shallow shelf). In: Hedgepth JW (ed) Treatise on marine ecology and paleoecology. Mem Geol Soc Amer 1
  3. 3,0 3,1 Sanders HL (1968) Marine benthic diversity: a comparative study. American Naturalist 102:243-282
  4. 4,0 4,1 Roy K, Jablonski D, Valentine JW, Rosenberg G (1998) Marine latitudinal diversity gradients; tests of casual hypotheses. Proc Natl Acad Sci USA 95:3699-3702
  5. Jablonski D, Roy K, Valentine J (2000) Analysing the latitudinal diversity gradient in marine bivalves. In: Harper EM, Taylor JD, Crame JA (eds) The evolutionary biology of the Bivalvia. p 361-365
  6. Clarke A, Lidgard S (2000) Spatial Patterns of Diversity in the Sea: Bryozoan Species Richness in the North Atlantic. Journal of Animal Ecology 69:799-814
  7. Culver SJ, Buzas MA (2000) Global latitudinal species diversity gradient in deep-sea benthic foraminifera. Deep-Sea Research 47:259-275
  8. Rex MA, Stuart CT, Coyne G (2000) Latitudinal gradients of species richness in the deep-sea benthos of North Atlantic. Proc. Natl Acad Sci USA 97:4082-4085
  9. Gray JS (2001) Marine diversity: the paradigms in patterns of species richness examined. Scientia Marina 65:41-56
  10. 10,0 10,1 Crame JA (2000) Evolution of taxonomic diversity gradients in the marine realm: evidence from the composition of Recent bivalve faunas. Paleobiology 26:188-214
  11. Clarke A (1992) Is there a latitudinal diversity cline in the sea? TREE 7:286-287
  12. Roy K, Jablonski D, Valentine JW (2000) Dissecting latitudinal diversity gradients: functional groups and clades of marine bivalves. Proc R Soc Lond B 267:293-299
  13. Rex MA, Stuart CT, Hessler RR, Allen JA, Sanders HL, Wilson GDF (1993) Global-scale latitudinal patterns of species diversity in the deep-sea benthos. Nature 365:636-639
  14. Kendall MA (1996) Are Arctic soft-sediment macrobenthic communities impoverished? Polar Biology 16:393-399
  15. Ellingsen KE, Gray JS (2002) Spatial patterns of benthic diversity: is there a latitudinal gradient along the Norwegian continental shelf? Journal of Animal Ecology 71:373-389
  16. Renaud and 34 others, in press, Continental-scale patterns in benthic invertebrate diversity: Insights from the MarBEF database, Mar Ecol Prog Ser.
  17. Kuklinski P, Barnes DKA, Taylor PDA (2006) Latitudinal patterns of diversity and abundance in North Atlantic intertidal boulder-fields. Mar Biol 147:1577–1583
  18. Grassle JF, Maciolek N (1992) Deep sea species richness: regional and local diversity estimates from quantitative bottom samples . American Naturalist 313-341
  19. Rex MA (1983) Geographic patterns of species diversity in the deep-sea benthos. In: Rowe GT (ed) The Sea vol 8. Deep Sea Biology. Wiley & sons, New York, p 453-472
  20. Rex MA, Etter RJ, Stuart C (1997) Large-scale patterns of species diversity in the deep-sea benthos . In: Ormond RJA, Gage JD, Angel MV (eds) Marine biodiversity: causes and consequences. Cambridge University Press, Cambridge, p 94-121
  21. Włodarska-Kowalczuk, M., Kendall, M.A., Węsławski, J.M., Klages, M., Soltwedel, T., (2004) Depth gradients of benthic standing stock and diversity on the continental margin at a high latitude ice-free site (off West Spitsbergen, 79 °N)., Deep-Sea Res I, 51, 1903-1914

The main author of this article is Włodarska- Kowalczuk, Maria
Please note that others may also have edited the contents of this article.