Temporal and spatial variability of environments drive the patterns of species richness along latitudinal, elevational, and depth gradients


This paper seeks to shed light on the primary causes of the latitudinal cline in species diversity, the asymmetry in species richness between the northern and southern hemispheres, and various patterns of species richness along mountain and continental slopes, which are at present of central interest in ecology. To that end, we restate, further develop and unify Janzen’s ideas about the higher fidelity of tropical organisms to their habitats; the notions of Sanders on temporal and spatial variations of environment and their impact on the breadth of species adaptations; the hypothesis of latitude-niche breadth and niche overlap; the theories of climatic stability, competitive exclusion and competitive divergence with the incorporation of some elements of the gradient model of diversification. We argue that during adaptation to a wide range of the same environmental factors in time, the high latitude species also become adapted to a wide range of those factors in space. As a result, they form not only very wide, but also widely overlapping ecological niches. This eventually leads to the competitive extinction of many species and a general impoverishment of biota. In contrast, relatively stable environments allow species to move more and more towards specialization with a simultaneous narrowing of their ecological niches that in turn leads to a reduction of niche overlap and greater species packing in communities. In tropical mountains and on the continental slope, where the environment is stable enough, the degree of its differentiation depends mainly on the steepness of slope. And since the steepest slopes tend to be located at intermediate elevations and intermediate bathyal depths, it is there that there are conditions for the highest specialization. These ideas can provide a framework for new approaches to biodiversity conservation of different regions.


competitive divergence, competitive extinction, environmental variability, niche overlap, specialization, species packing, species richness


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Acharya, B. K., Sanders, N. J., Vijayan, L., and Chettri, B. 2011. Elevational gradients in bird diversity in the Eastern Himalaya: an evaluation of distribution patterns and their underlying mechanisms. PLoS ONE 6:e29097. https://doi.org/10.1371/journal.pone.0029097

Addo-Bediako, A. S., Chown, S. L., and Gaston, K. J. 2000. Thermal tolerance, climatic variability and latitude. Proceedings of the Royal Society B 267:739−745. https://doi.org/10.1098/rspb.2000.1065

Adler, P. B., Seabloom, E. W., Borer, E. T., Hillebrand, H., …, and Yang, L. H. 2011. Productivity is a poor predictor of plant species richness. Science 333:1750−1753. https://doi.org/10.1126/science.1204498

Angel, M. A. 1993. Biodiversity of the pelagic ocean. Conservation Biology 7:760−772. https://doi.org/10.1046/j.1523-1739.1993.740760.x

Archibald, S. B., Bossert, W. H., Greenwood, D. R., and Farrell, B. D. 2010. Seasonality, the latitudinal gradient of diversity, and Eocene insects. Paleobiology 36:374−398. https://doi.org/10.1016/j.palaeo.2012.10.043

Barthlott, W., Mutke, J., Rafiqpoor, D., Kier, G., and Kreft, H. 2005. Global centers of vascular plant diversity. Nova Acta Leopoldina 92:61−83.

Billet, D. S. M., Lampitt, R. S., Rice, A. L., and Mantoura, R. F. C. 1983. Seasonal sedimentation of phytoplankton to the deep-sea benthos. Nature 302:520−522. https://doi.org/10.1038/302520a0

Blackburn, T. M., Gaston, K. J. 1996. Spatial patterns in the species richness of birds in the New World. Ecography 19:369−376. https://doi.org/10.1111/j.1600-0587.1996.tb00001.x

Boltovskoy, D., Gibbons, M. J., Hutchings, L., and Binet, D. 1999. General biological features of the South Atlantic; pp. 1−42 in Boltovskoy, D. (Ed) South Atlantic zooplankton, Leiden, Backhuys Publishers.

Boucher, G., and Lambshead, P. J. D. 1995. Ecological biodiversity of marine nematodes in samples from temperate, tropical, and deep-sea regions. Conservation Biology 9:1594−1604. https://doi.org/10.1046/j.1523-1739.1995.09061594.x

Brown, W. L. Jr., and Wilson, E. O. 1956. Character displacement. Systematic Zoology 5:49−64. https://doi.org/10.2307/2411924

Buckley, L. B., and Jetz, W. 2007. Environmental and historical constraints on global patterns of amphibian richness. Proceedings of the Royal Society B 274:1167−1173. https://doi.org/10.1098/rspb.2006.0436

Bukvareva, E., and Aleshchenko, G. 2013. Optimization, niche and neutral mechanisms in the formation of biodiversity. American Journal of Life Sciences 1(4):174−183. https://doi.org/10.11648/j.ajls.20130104.16

Cadena, C. D., Kozak, K. H., Gómez, J. P., Parra, J. L., …, and Graham, C. H. 2011. Latitude, elevational climatic zonation and speciation in New World vertebrates. Proceedings of the Royal Society B 279:194−201. https://doi.org/10.1098/rspb.2011.0720

Chettri, B., Bhupathy, S., and Acharya, B. K. 2010. Distribution pattern of reptiles along an eastern Himalayan elevation gradient, India. Acta Oecologica 36:16−22. https://doi.org/10.1016/j.actao.2009.09.004

Clarke, A., and Gaston, K. J. 2006. Climate, energy and diversity. Proceedings of the Royal Society B 273:2257−2266. https://doi.org/10.1098/rspb.2006.3545

Colwell, M. A. 2010. Shorebird ecology, conservation, and management. 344 pp. University of California Press, Berkeley and Los Angeles.

Cousins, S. H. 1989. Species richness and the energy theory. Nature 340:350−351. https://doi.org/10.1038/340350b0

Crame, J. A. 2000. The nature and origin of taxonomic diversity gradients in marine bivalves; pp. 347−360 in: Harper E. M., Taylor J. D., and Crame J. A. (Eds) The evolutionary biology of the Bivalvia. Geological Society of London, Special Publication, 177, London, Alden Press.

Currie, D. J. 1991. Energy and large-scale patterns of animal- and plant-species richness. American Naturalist 137:27−49. https://doi.org/10.1086/285144

Darwin, C. 1845/2008. The Voyage of the Beagle, Cosimo, New York.

Darwin, C. 1872. The Origin of Species by Means of Natural Selection or the Preservation of Favoured Races in the Struggle for Life, 6th edition, John Murray, London.

Davies, R. G., Orme, C. D. L., Storch, D., Olson, V. A., …, and Gaston, K. J. 2007. Topography, energy and the global distribution of bird species richness. Proceedings of the Royal Society B 274:1189−1197. https://doi.org/10.1098/rspb.2007.0046

de Klerk, H. M., Crowe, T. M., Fjeldså, J., and Burgess, N. D. 2002. Biogeographical patterns of endemic terrestrial Afrotropical birds. Diversity and Distributions 8:147−162. https://doi.org/10.1046/j.1472-4642.2002.00142.x

de Oliveira, T. G. 1994. Neotropical cats: ecology and conservation. São Luís, Edufma.

Dobzhansky, Th. 1950. Evolution in the tropics. American Scientist 38:209−221.

Dodge, J. D., and Marshall, H. G. 1994. Biogeographic analysis of the armored planktonic dinoflagellate Ceratium in the North Atlantic and adjacent seas. Journal of Phycology 30:905−922. https://doi.org/10.1111/j.0022-3646.1994.00905.x

Dyer, L. A., Singer, M. S., Lill, J. T., Stireman, J. O., …, and Coley, P. D. 2007. Host specificity of Lepidoptera in tropical and temperate forests. Nature 448:696−700. https://doi.org/10.1038/nature05884

Eggleton, P. 1994. Termites live in a pear-shaped world: a response to Platnick. Journal of Natural History 28:1209−1212. https://doi.org/10.1080/00222939400770611

Emlin, J. M. 1966. The role of time and energy in food preferences. American Naturalist 100:611−617. https://doi.org/10.1086/282455

Evans, K. L., Warren, P. H., Gaston, K. J. 2005. Species-energy relationships at the macroecological scale: a review of the mechanisms. Biological Reviews 80:1−25. https://doi.org/10.1017/S1464793104006517

Fischer, A. G. 1960. Latitudinal variation in organic diversity. Evolution 14:64−81. https://doi.org/10.1111/j.1558-5646.1960.tb03057.x

Fjeldså, J., and Rahbek, C. 2006. Diversification of tanagers, a species-rich bird group, from the lowlands to montane regions of South America. Integrative and Comparative Biology 46:72−81. https://doi.org/10.1093/icb/icj009

Fjeldså, J., Kiure, J., Doggart, N., Hansen, L. A., and Perkin, A. 2010. Distribution of highland forest birds across a potential dispersal barrier in the Eastern Arc Mountains of Tanzania. Steenstrupia 32:1−43.

Floeter, S. R., and Soares-Gomes, A. 1999. Biogeographic and species richness patterns of Gastropoda on the southwestern Atlantic. Revista Brasileira de Biologia 59:567−575. https://doi.org/10.1590/S0034-71081999000400006

France, R. 1992. The North American latitudinal gradient in species richness and geographical range of freshwater crayfish and amphipods. American Naturalist 139:342−354. https://doi.org/10.1086/285330

Fuhrman, J. A., Steele, J. A., Hewson, I., Schwalbach, M. S., …, and Brown, J. H. 2008. A latitudinal diversity gradient in planktonic marine bacteria. Proceedings of the National Academy of Sciences USA 105:7774−7778. https://doi.org/10.1073/pnas.0803070105

Gardner, A. L. 2007. Mammals of South America. Volume 1: Marsupials, Xenarthrans, Shrews and Bats. University of Chicago Press, Chicago.

Gaston, A. J., and Jones, I. L. 1998. The auks: Alcidae. Oxford University Press, Oxford.

Gaston, K. J., and Chown, S. L. 1999. Why Rapoport’s rule does not generalise. Oikos 84:309−312. https://doi.org/10.2307/3546727

Gaston K. J. 2000. Global patterns in biodiversity. Nature 405:220−227. https://doi.org/10.1038/35012228

Gause, G. F. 1934. The Struggle for Existence. Williams and Wilkins, Baltimore.

Ghalambor, C. K., Huey, R. B., Martin, P. R., Tewksbury, J. J., and Wang, G. 2006. Are mountain passes higher in the tropics? Janzen’s hypothesis revisited. Integrative and Comparative Biology 46:5−17. https://doi.org/10.1093/icb/icj003

Gillman, L. N., and Wright, S. D. 2006. The influence of productivity on the species richness of plants: a critical assessment. Ecology 87:1234−1243. https://doi.org/10.1890/0012-9658(2006)87[1234:TIOPOT]2.0.CO;2

Goodman, S. M., and Rasolonandrasana, B. P. N. 2001. Elevational zonation of birds, insectivores, rodents and primates on the slopes of the Andringitra Massif, Madagascar. Journal of Natural History 35:285−305. https://doi.org/10.1080/00222930150215387

Graham, G. L. 1990. Bats versus birds: comparisons among Peruvian volant vertebrate faunas along an elevational gradient. Journal of Biogeography 17:657−668. https://doi.org/10.2307/2845147

Grant, P. R. 1972. Convergent and divergent character displacement. Biological Journal of the Linnean Society 4:39−68. https://doi.org/10.1111/j.1095-8312.1972.tb00690.x

Grinnell, J., and Storer, T. I. 1924. Animal Life in the Yosemite. University of California Press, Berkeley.

Haedrich, R. L., Rowe, G. T., and Polloni, P. T. 1980. The megabenthic fauna in the deep sea south of New England, USA. Marine Biology 57:165−179. https://doi.org/10.1007/BF00390735

Hall, J. P. W. 2005. Montane speciation patterns in Ithomiola butterflies (Lepidoptera: Riodinidae): are they consistently moving up in the world? Proceedings of the Royal Society B 272:2457−2466. https://doi.org/10.1098/rspb.2005.3254

Hartmann, D. L. 1994. Global physical climatology. Academic Press, San Diego.

Heaney, L. R. 2001. Small mammal diversity along elevational gradients in the Philippines: an assessment of patterns and hypotheses. Global Ecology and Biogeography 10:15−39. https://doi.org/10.1046/j.1466-822x.2001.00227.x

Herzog, S. K., Kessler, M., and Bach, K. 2005. The elevational gradient in Andean bird species richness at the local scale: a foothill peak and a high-elevation plateau. Ecography 28:209−222. https://doi.org/10.1111/j.0906-7590.2005.03935.x

Hessler, R. R., and Sanders, H. L. 1967. Faunal diversity in the deep-sea. Deep Sea Research and Oceanographic 14:65−78. https://doi.org/10.1016/0011-7471(67)90029-0

Hoare, B. 2009. Animal migration: remarkable journeys in the wild. University of California Press, Berkeley.

Hua, X., and Wiens, J. J. 2010. Latitudinal variation in speciation mechanisms in frogs. Evolution 64:429−443. https://doi.org/10.1111/j.1558-5646.2009.00836.x

Huston, M. A. 1979. A general hypothesis of species diversity. American Naturalist 113:81−101. https://doi.org/10.1086/283366

Huston, M. A. 1985a. Patterns of species diversity in relation to depth at Discovery Bay, Jamaica. Bulletin of Marine Science 37:928−935.

Huston, M. A. 1985b. Patterns of species diversity on coral reef. Annual Review of Ecology and Systematics 16:149−177. https://doi.org/10.1146/annurev.es.16.110185.001053

Huston, M. A. 1994. Biological diversity. Cambridge University Press, Cambridge.

Hutchinson, G. E. 1959. Homage to Santa Rosalia, or why are there so many kinds of animals? American Naturalist 93:145−159. https://doi.org/10.1086/282070

Janzen, D. H. 1967. Why mountain passes are higher in the tropics. American Naturalist 101:233−249. https://doi.org/10.1086/282487

Janzen, D. H. 1981. The peak in North American ichneumonid species richness lies between 38o and 42o N. Ecology 62:532−537. https://doi.org/10.2307/1937717

Johnsgard, P. A. 1981. The plovers, sandpipers, and snipes of the world, University of Nebraska Press, Lincoln.

Jones, N. S., and Sanders, H. L. 1972. Distribution of Cumacea in the deep Atlantic. Deep Sea Research and Oceanographic Abstracts 19:737−745. https://doi.org/10.1016/0011-7471(72)90066-6

Jonzen, N., Knudsen, E., Holt, R., Saether, B.-E. 2011. In: Milner-Gulland, E. J., Fryxell, J. M., and Sinclair, A. R. E. (Eds) Animal migration: a synthesis, Oxford, Oxford University Press 91−109.

Kolomiytsev, N. P., and Poddubnaya, N. Ya. 2007. The origin of life as a result of changing the evolutionary mechanism. Rivista di Biologia / Biology Forum 100:11−16.

Klopfer, P. H. 1959. Environmental determinants of faunal diversity. American Naturalist 93:337−342. https://doi.org/10.1086/282092

Klopfer, P. H., and MacArthur, R. 1961. On the causes of tropical species diversity: niche overlap. American Naturalist 95:223−226. https://doi.org/10.1086/282179

Kolasa, J., Hewitt, C. L., and Drake, J. A. 1998. Rapoport’s rule: an explanation or a byproduct of the latitudinal gradient in species richness? Biodiversity and Conservation 7:1447−1455. https://doi.org/10.1023/A:1008805230673

Kozak, K. H., and Wiens, J. J. 2006. Does niche conservatism promote speciation? A case study in North American salamanders. Evolution 60:2604−2621. https://doi.org/10.1554/06-334.1

Kozak, K. H., and Wiens, J. J. 2007. Climatic zonation drives latitudinal variation in speciation mechanisms. Proceedings of the Royal Society B 274:2995−3003. https://doi.org/10.1098/rspb.2007.1106

Kozak, K. H., and Wiens, J. J. 2010. Accelerated rates of climatic-niche evolution underlie rapid species diversification. Ecology Letters 13:1378−1389. https://doi.org/10.1111/j.1461-0248.2010.01530.x

Küper, W., Kreft, H., Nieder, J., Köster, N., and Barthlott, W. 2004. Large-scale diversity patterns of vascular epiphytes in Neotropical montane rain forests. Journal of Biogeography 31:1477−1487. https://doi.org/10.1111/j.1365-2699.2004.01093.x

Larsen, T. H., Escobar, F., and Armbrecht, I. 2011. Insects of the Tropical Andes: Diversity Patterns, Processes and Global Change; pp. 228−244 in: Herzog, S. K., Martínez, R., Joergensen, P. M., and Tiessen, H. (Eds). Climate change effects on the biodiversity of the tropical Andes: an assessment of the status of scientific knowledge. Brazil and Paris, Inter-American Institute for Global Change Research and Scientific Committee on Problems of the Environment, São José dos Campos.

Letcher, A. J., and Harvey, P. H. 1994. Variation in geographical range size among mammals of the Palearctic. American Naturalist 144:30−42. https://doi.org/10.1086/285659

Levin, L. A. 2003. Oxygen minimum zone benthos: adaptation and community response to hypoxia. Oceanography and Marine Biology: an Annual Review 41:1−45.

Li, J., He, Q., Hua, X., Zhou, J., Xu, H., Chen, J., and Fu, C. 2009. Climate and history explain the species richness peak at mid-elevation for Schizothorax fishes (Cypriniformes: Cyprinidae) distributed in the Tibetan Plateau and its adjacent regions. Global Ecology and Biogeography 18:123−136. https://doi.org/10.1111/j.1466-8238.2008.00430.x

Lomolino, M. V., Riddle, B. R., Whittaker, R. J. and Brown, J. H. 2010. Biogeography, 4th edition.

Luo, Z., Tang, S., Li, C., Fang, H., Hu, H., Yang, J., Ding, J., and Jiang, Z. 2012. Environmental effects on vertebrate species richness: testing the energy, environmental stability and habitat heterogeneity hypotheses. PLoS ONE 7:e35514. https://doi.org/10.1371/journal.pone.0035514

MacArthur, J. W. 1975 Environmental fluctuations and species diversity; pp. 74−80 in Cody, ML & Diamond, JM (Eds) Ecology and evolution of communities. Cambridge, Belknap Press.

MacArthur, R. H. 1955. Fluctuations of animal populations and a measure of community stability. Ecology 36:533−536. https://doi.org/10.2307/1929601

MacArthur, R. H. 1972. Geographical ecology: patterns in the distribution of species. Princeton University Press, Princeton.

MacArthur, R. H. and MacArthur, J. W. 1961. On bird species diversity. Ecology 42:594−598. https://doi.org/10.2307/1932254

MacArthur, R. H. and Wilson, E. O. 1967. The theory of island biogeography. Princeton University Press, Princeton.

Macpherson, E. 2002. Large-scale species-richness gradients in the Atlantic Ocean. Proceedings of the Royal Society B 95:3699−3702. https://doi.org/10.1098/rspb.2002.2091

Mani, M. S. 1968. Ecology and biogeography of high altitude insects. Series Entomologica. Vol. 4. Dr. W. Junk N. V. Publishers, The Hague.

Martin, P. R. and McKay, J. K. 2004. Latitudinal variation in genetic divergence of populations and the potential for future speciation. Evolution 58:938−945. https://doi.org/10.1111/j.0014-3820.2004.tb00428.x

Martin, P. R. and Tewksbury, J. J. 2008. Latitudinal variation in subspecific diversification of birds. Evolution 62:2775−2788. https://doi.org/10.1111/j.1558-5646.2008.00489.x

Mayr, E. 1963. Animal species and evolution. Harvard University Press, Cambridge.

McCain, C. M. 2009. Vertebrate range sizes indicate that mountains may be ‘higher’ in the tropics. Ecology Letters 12:550−560. https://doi.org/10.1111/j.1461-0248.2009.01308.x

McCain, C. M. and Grytnes, J.-A. 2010. Elevational gradients in species richness; pp. 1−10 in Encyclopedia of Life Sciences (ELS), Chichester, John Wiley and Sons (Ltd). https://doi.org/10.1002/9780470015902.a0022548

Menzies, R. J., George, R. Y., and Rowe, G. T. 1973. Abyssal environment and ecology of the world oceans. Wiley-Interscience, New York and London.

Mittelbach, G. G., Steiner, C. F., Scheiner, S. M., Gross, K. L., … Gough, L. 2001. What is the observed relationship between species richness and productivity? Ecology 82:2381−2396. https://doi.org/10.1890/0012-9658(2001)082[2381:WITORB]2.0.CO;2

Mittelbach, G. G., Schemske, D. W., Cornell, H. V., Allen, A. P., … Turelli, M. 2007. Evolution and the latitudinal diversity gradient: speciation, extinction and biogeography. Ecology Letters 10:315−331. https://doi.org/10.1111/j.1461-0248.2007.01020.x

Morse, D. H. 1971. The insectivorous bird as an adaptive strategy. Annual Review of Ecology and Systematics 2:177−200. https://doi.org/10.1146/annurev.es.02.110171.001141

Newton, I. 2008. The Migration Ecology of Birds. Academic Press, London.

Nor, S. M. D. 2001. Elevational diversity patterns of small mammals on Mount Kinabalu, Sabah, Malaysia. Global Ecology and Biogeography 10:41−62. https://doi.org/10.1046/j.1466-822x.2001.00231.x

Nybakken, J. W., Bertness, M. D. 2004. Marine Biology: An Ecological Approach. 6th ed, Benjamin Cummings, San Francisco.

O’Brien, E. M. 1993. Climatic gradients in woody plant species richness: towards an explanation based on an analysis of southern Africa’s woody flora. Journal of Biogeography 20:181−198. https://doi.org/10.2307/2845670

Ogden, R. and Thorpe, R. S. 2002. Molecular evidence for ecological speciation in tropical habitats. Proceedings of the National Academy of Sciences USA 99:13612−13615. https://doi.org/10.1073/pnas.212248499

Olabarria, C. 2006. Faunal change and bathymetric diversity gradient in deep-sea prosobranchs from Northeastern Atlantic. Biodiversity and Conservation 15:3685−3702. https://doi.org/10.1007/s10531-005-1344-9

Ollerton, J. and Cranmer, L. 2002. Latitudinal trends in plantpollinator interactions: Are tropical plants more specialized? Oikos 98:340−350. https://doi.org/10.1034/j.1600-0706.2002.980215.x

Oömmen, M. A. and Shanker, K. 2005. Elevational species richness patterns emerge from multiple local scale mechanisms in Himalayan plants. Ecology 86:3039−3047. https://doi.org/10.1890/04-1837

Orme, C. D., Davies, R. G., Olson, V. A., …, and Gaston, K. J. 2006. Global patterns of geographic range size in birds. PLoS Biology 4:e208. https://doi.org/10.1371/journal.pbio.0040208

Owen, R. W. 1981. Fronts and eddies in the sea: mechanisms, interactions and biological effects; pp. 197−233 in: Longhurst, AR (Ed) Analysis of marine ecosystems. London, Academic Press.

Pärtel, M., Laanisto, L., and Zobel, M. 2007. Contrasting plant productivity–diversity relationships across latitude: the role of evolutionary history. Ecology 88:1091−1097. https://doi.org/10.1890/06-0997

Peck, S. B. and Kaulbars, M. M. 1987. A synopsis of the distribution of the carrion beetles (Coleoptera: Silphidae) of the conterminous United States. Proceedings of the Entomological Society of Ontario 188:47−81.

Pianka, E. R. 1966. Latitudinal gradients in species diversity: a review of the concepts. American Naturalist 100:33−46. https://doi.org/10.1086/282398

Pianka, E. R. 1970. On r- and K-selection. American Naturalist 104:592−597. https://doi.org/10.1086/282697

Pielou, E. C. 1979. Biogeography. Wiley Interscience, New York.

Platnick, N. I. 1991. Patterns of biodiversity: tropical vs temperate. Journal of Natural History 25:1083−1088. https://doi.org/10.1080/00222939100770701

Powney, G. D., Grenyer, R., Orme, C. D. L., Owens, I. P. F., and Meiri, S. 2010. Hot, dry and different: Australian lizard richness is unlike that of mammals, amphibians and birds. Global Ecology and Biogeography 19:386−396. https://doi.org/10.1111/j.1466-8238.2009.00521.x

Pringle, C. M. 2000. River conservation in Latin America and the Caribbean; pp. 371−384 in Boon, PJ, Davies, BR & Petts, GE (Eds) Global Perspectives on River Conservation: Science, Policy and Practice. Chichester, John Wiley and Sons.

Pyron, R. A. and Burbrink, F. T. 2009. Can the tropical conservatism hypothesis explain temperate species richness patterns? An inverse latitudinal biodiversity gradient in the New World snake tribe Lampropeltini. Global Ecology and Biogeography 18:406−415. https://doi.org/10.1111/j.1466-8238.2009.00462.x

Rabosky, D. L., Chang, J., Title, P. O., Cowman, P. F., Sallan, L., Friedman, M., Kaschner, K., Garilao, C., Near, Th. J., Coll, M., and Alfaro, M. E. 2018. An inverse latitudinal gradient in speciation rate for marine fishes. Nature 559:392–395. https://doi.org/10.1038/s41586-018-0273-1

Rahbek, C. 1995. The elevational gradient of species richness: a uniform pattern? Ecography 18:200−205. https://doi.org/10.1111/j.1600-0587.1995.tb00341.x

Rahbek, C. and Graves, G. R. 2001. Multiscale assessment of patterns of avian species richness. Proceedings of the National Academy of Sciences USA 98:4534−4539. https://doi.org/10.1073/pnas.071034898

Rapoport, E. H. 1982. Areography: Geographical Strategies of Species, Pergamon Press, Oxford.

Renjifo, L. M., Servat, G. P., Goerck, J. M., Loiselle, B. A., and Blake, J. G. 1997. Patterns of species composition and endemism in the northern Neotropics: a case for conservation of montane avifaunas. Ornithological Monographs 48:577−594. https://doi.org/10.2307/40157554

Rex, M. A. 1973. Deep-sea species diversity: decreased gastropod diversity at abyssal depths. Science 181:1051−1053. https://doi.org/10.1126/science.181.4104.1051

Rex, M. A. 1981. Community structure in the deep-sea benthos. Annual Review of Ecology and Systematics 12:331−353. https://doi.org/10.1146/annurev.es.12.110181.001555

Rex, M. A. and Etter, R. J. 2010. Deep-Sea Biodiversity: Pattern and Scale. Harvard University Press, Cambridge.

Rex, M. A., Stuart, C. T., and Coyne, G. 2000. Latitudinal gradients of species richness in the deep-sea benthos of the North Atlantic. Proceedings of the National Academy of Sciences USA 97:4082−4085. https://doi.org/10.1073/pnas.050589497

Ricklefs, R. E. and Latham, R. E. 1992. Intercontinental correlation of geographical ranges suggests stasis in ecological traits of relict genera of temperate perennial herbs. American Naturalist 139:1305−1321. https://doi.org/10.1086/285388

Rosenzweig, M. L. 1995. Species diversity in space and time, Cambridge University Press, Cambridge.

Roy, M. S. 1997. Recent diversification of African greenbuls (Pycnonotidae: Andropadus) supports a montane speciation model. Proceedings of the Royal Society B 264:1337−1344. https://doi.org/10.1098/rspb.1997.0185

Roy, K., Jablonski, D., Valentine, J. W., and Rosenberg, G. 1998. Marine latitudinal diversity gradients: tests of causal hypotheses. Proceedings of the National Academy of Sciences USA 95:3699−3702. https://doi.org/10.1073/pnas.95.7.3699

Ruggiero, A. 1994. Latitudinal correlates of the sizes of mammalian geographical ranges in South America. Journal of Biogeography 21:545−559. https://doi.org/10.2307/2845657

Rutherford, S., D’Hondt, S., and Prell, W. 1999. Environmental controls on the geographic distribution of zooplankton diversity. Nature 400:749−753. https://doi.org/10.1038/23449

Sanders, H. L. 1968. Marine benthic diversity: a comparative study. American Naturalist 102:243−282. https://doi.org/10.1086/282541

Sarmiento, G. 1986. Ecologically crucial features of climate in high tropical mountains; pp. 11−46 in Vuilleumier, F. and Monasterio, M. (Eds). High altitude tropical biogeography. New York, Oxford University Press.

Schmalhausen, I. I. 1949. Factors of Evolution: The Theory of Stabilizing Selection. Blakiston, Philadelphia.

Schmit, J. P. 2005. Species richness of tropical wood-inhabiting macrofungi provides support for species-energy theory. Mycologia 97:751−761. https://doi.org/10.1080/15572536.2006.11832766

Scriber, J. M. 1973. Latitudinal gradients in larval feeding specialization of the world Papilionidae (Lepidoptera). Psyche 80:355−373. https://doi.org/10.1155/1973/52610

Sheppard, C. R. C. 1980. Coral cover, zonation and diversity on reef slopes of Chagos Atolls, and population structures of the major species. Marine Ecology — Progress Series 2:193−205. https://doi.org/10.3354/meps002193

Slansky, F. 1972. Latitudinal gradients in species diversity of the New World swallowtail butterflies. Journal of Research on the Lepidoptera 11:201−218.

Slobodkin, L. B. and Sanders, H. L. 1969. On the contribution of environmental predictability to species diversity. Brookhaven Symposium in Biology 22:82−95.

Slove, J. and Janz, N. 2010. Phylogenetic analysis of the latitude-niche breadth hypothesis in the butterfly subfamily Nymphalinae. Ecological Entomology 35:768−774. https://doi.org/10.1111/j.1365-2311.2010.01238.x

Smith, S. A., de Oca, A. N. M., Reeder, T. W., and Wiens, T. W. 2007. A phylogenetic perspective on elevational species richness patterns in Middle American treefrogs: why so few species in lowland tropical rainforests? Evolution 61:1188–1207. https://doi.org/10.1111/j.1558-5646.2007.00085.x

Stevens, G. C. 1989. The latitudinal gradient in geographical range: how so many species coexist in the tropics. American Naturalist 133:240−256. https://doi.org/10.1086/284913

Stuart, C. T., Rex, M. A., and Etter, R. J. 2003. Large−scale spatial and temporal patterns of deap−sea benthic species diversity; pp. 297−313 in Tyler, P. A. (Ed) Ecosystems of the world. Volume 28. Ecosystems of the deep oceans. Amsterdam, Elsevier.

Turner, J. R. G., Gatehouse, C. M., and Corey, C. A. 1987. Does solar energy control organic diversity? Butterflies, moths and the British climate. Oikos 48:195−205. https://doi.org/10.2307/3565855

Väinölä, R., Witt, J. D. S., Grabowski, M., Bradbury, J. H., Jazdzewski, K., and Sket, B. 2008. Global diversity of amphipods (Amphipoda; Crustacea) in freshwater. Hydrobiologia 595:241−255. https://doi.org/10.1007/s10750-007-9020-6

Valdovinos, C., Navarette, S. A., and Marquet, P. A. 2003. Mollusk species diversity in the Southeastern Pacific: why are there more species towards the pole? Ecography 26:139−144. https://doi.org/10.1034/j.1600-0587.2003.03349.x

Veron, J. E. N. 1995. Corals in space and time: The biogeography and evolution of the scleractinia. University of the New South Wales Press, New South Wales, Sydney.

von Humboldt, A. 1808. Ansichten der Natur: mit wissenschaftlichen Erläuterungen. J. G. Cotta‘schen Buchhandlung, Tübingen.

Wallace, A. R. 1878. Tropical nature and other essays. Macmillan, New York.

Wiens, J. J. 2007. Global patterns of species richness and diversification in amphibians. American Naturalist 170:86−106. https://doi.org/10.1086/519396

Williams, C. B. 1964. Patterns in the balance of nature: and related problems in quantitative ecology. Academic Press, New York.

Williams, T. D. 1995. The penguins: Spheniscidae. Oxford University Press, Oxford.

Williams, Y. M., Williams, S. E., Alford, R. A., Waycott, M., and Johnson, C. N. 2006. Niche breadth and geographical range: ecological compensation for geographical rarity in rainforest frogs. Biology Letters 2:532−535. https://doi.org/10.1098/rsbl.2006.0541

Willig, M. R., Kaufman, D. M., and Stevens, R. D. 2003. Latitudinal gradients of biodiversity: pattern, process, scale, and synthesis. Annual Review of Ecology, Evolution, and Systematics 34:273−309. https://doi.org/10.1146/annurev.ecolsys.34.012103.144032

Wilson, D. E, and Reeder, D. M. (Eds) 2005. Mammal Species of the World: A Taxonomic and Geographic Reference, 3rd ed, Baltimor, Johns Hopkins University Press.

Woodd-Walker, R. S., Ward, P., and Clarke, A. 2002. Large scale patterns in diversity and community structure of surface water copepods from the Atlantic Ocean. Marine Ecology Progress Series 236:189−203. https://doi.org/10.3354/meps236189

Wright, D. H., Currie, D. J., and Maurer, B. A. 1993. Energy supply and patterns of species richness on local and regional scales; pp. 66−74 in Ricklefs, R. E. and Schluter D. (Eds) Species Diversity in Ecological Communities. Chicago, University of Chicago Press.

Wright, S. J. 2002. Plant diversity in tropical forests: a review of mechanisms of species coexistence. Oecologia 130:1–14. https://doi.org/10.1007/s004420100809

Wu, Y., Colwell, R. K., Rahbek, C., Zhang, C., Quan, Q., Wang, C., and Le, L. 2013. Explaining the species richness of birds along a subtropical elevational gradient in the Hengduan Mountains. Journal of Biogeography 40:2310−2323. https://doi.org/10.1111/jbi.12177
How to Cite
Kolomiytsev, N., & Poddubnaya, N. (2018). Temporal and spatial variability of environments drive the patterns of species richness along latitudinal, elevational, and depth gradients. Biological Communications, 63(3), 189–201. https://doi.org/10.21638/spbu03.2018.305
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