Abstract
Size-latitude trends in the meiobenthic phylum Kinorhyncha, commonly known as mud dragons, have been explored in oceans worldwide. Generalized least squares regression was used to assess relationships between size and latitude, as well as between size, latitude, and two selected environmental variables that exhibit latitudinal gradation: the sea surface temperature and the net primary productivity. Different structures of spatial autocorrelation and potential confounding factors, such as the species richness and the number of kinorhynch records that could affect latitudinal gradients, were also addressed. In addition, generalized mixed models were used to determine the influence of the phylogeny on body size. Size-latitude relationships of Kinorhyncha were commonly found globally, as well as for particular geographic regions (hemispheres and/or coastlines), with important differences between taxonomic groups. These size-latitude trends were heterogeneous and implied the influence of the latitude itself, environmental variables, and phylogeny. These facts indicate that a single underlying process is not likely to explain the observed relationships but a complex interaction of several macroecological patterns both present and past. Perhaps, the inclusion of future new reports, conducted in undersampled areas, may shed some light on the matter and reveal more generalized size-latitude patterns. Nevertheless, it is also likely that broadly generalizable size-latitude relationships may not exist in meiofaunal communities.
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Data availability
The original data used in this study are available within the article (Supporting Information Appendix S1). Data on environmental variables used in the present study are available in the following public domains: US National Oceanic and Atmospheric Administration (NOAA) (http://noaa.gov/) and Guillaume Maze’s data site (http://data.guillaumemaze.org/). Geographic occurrences of all the species were extracted from the literature, which are referenced in the Supporting Information Appendix S1.
References
Adrianov, A. V., & Maiorova, A. S. (2019). Echinoderes ultraabyssalis sp. nov. from the Kuril-Kamchatka Trench – the first hadal representative of the Kinorhyncha (Kinorhyncha: Cyclorhagida). Progress in Oceanography, 178, e102142. https://doi.org/10.1016/j.pocean.2019.102142.
Alho, J. S., Herczeg, G., Laugen, A. T., Räsänen, K., Laurila, A., & Merilä, J. (2010). Allen’s rule revisited: quantitative genetics of extremity length in the common frog along a latitudinal gradient. Journal of Evolutionary Biology, 24(1), 59–70. https://doi.org/10.1111/j.1420-9101.2010.02141.x.
Allen, J. A. (1877). The influence of physical conditions in the genesis of species. Radical Review, 1, 108–140.
Arendt, J. D. (2011). Size-fecundity relationships, growth trajectories, and the temperature-size rule for ectotherms. Evolution, 65, 43–51. https://doi.org/10.1111/j.1558-5646.2010.01112.x.
Arnett, A. E., & Gotelli, N. J. (2003). Bergmann’s rule in larval ant lions: testing the starvation resistance hypothesis. Ecological Entomology, 28, 645–650. https://doi.org/10.1111/j.1365-2311.2003.00554.x.
Armenteros, M., & Ruiz-Abierno, A. (2015). Body size distribution of free-living marine nematodes from a Caribbean coral reef. Nematology, 17, 1153–1164 https://doi.org/10.1163/15685411-00002930.
Ashton, K. G., & Feldman, C. R. (2003). Bergmann’s rule in non-avian reptiles: turtles follow it, lizards and snakes reverse it. Evolution, 57, 1151–1163. https://doi.org/10.1111/j.0014-3820.2003.tb00324.x.
Ashton-Acton, Q. (2013). Issues in global environment – globalization and global change research (3rd edn.). Atlanta: ScholarlyEditions.
Atkinson, D. (1994). Temperature and organism size: a biological law of ectotherms? Advances in Ecological Research, 25, 1–58. https://doi.org/10.1016/S0065-2504(08)60212-3.
Atkinson, D. (1995). Effects of temperature on the size of aquatic ectotherms: exceptions to the general rule. Journal of Thermal Biology, 20(1/2), 61–74. https://doi.org/10.1016/0306-4565(94)00028-H.
Azovsky, A., & Mazei, Y. (2013). Large-scale patterns in the diversity and distribution of marine benthic ciliates: do microbes have macroecology? Global Ecology and Biogeography, 22, 163–172. https://doi.org/10.1111/j.1466-8238.2012.00776.x.
Barnes, R. S. K. (2010). Regional and latitudinal variation in the diversity, dominance and abundance of microphagous microgastropods and other benthos in intertidal beds of dwarf eelgrass, Nanozostera spp. Marine Biodiversity, 40, 95–106. https://doi.org/10.1007/s12526-010-0036-1.
Bartels, P. J., Fontaneto, D., Roszkowska, M., Nelson, D. R., & Kaczmarek, Ł. (2019). Latitudinal gradients in body size in marine tardigrades. Zoological Journal of the Linnean Society. https://doi.org/10.1093/zoolinnean/zlz080.
Belk, M. C., & Houston, D. D. (2002). Notes and comments Bergmann’s rule in ectotherms: a test using freshwater fishes. American Naturalist, 160(6), 83–88. https://doi.org/10.1086/343880.
Berggreen, U., Hansen, B., & Kiørboe, T. (1988). Food size spectra, ingestion and growth of the copepod Acartia tonsa during development: implications for determination of copepod production. Marine Biology, 99, 341–352. https://doi.org/10.1007/BF02112126.
Bergmann, C. (1848). Über die Verhältnisse der Wärmeökonomie der Thiere zu ihrer Grösse. Göttinger Studien, 3, 595–708.
Berke, S. K., Jablonski, D., Krug, A. Z., Roy, K., & Tomasovych, A. (2013). Beyond Bergmann’s rule: size-latitude relationships in marine Bivalvia world-wide. Global Ecology and Biogeography, 22, 173–183. https://doi.org/10.1111/j.1466-8238.2012.00775.x.
Blackburn, T. M., Gaston, K. J., & Loder, N. (1999). Geographic gradients in body size: a clarification of Bergmann’s rule. Diversity and Distributions, 5, 165–174. https://doi.org/10.1046/j.1472-4642.1999.00046.x.
Blackburn, T. M., & Hawkins, B. A. (2004). Bergmann’s rule and the mammal fauna of northern North America. Ecography, 27, 715–724. https://doi.org/10.1111/j.0906-7590.2004.03999.x.
Blanckenhorn, W. U., & Demont, M. (2004). Bergmann and converse Bergmann latitudinal clines in arthropods: two ends of a continuum? Integrative and Comparative Biology, 44, 413–424. https://doi.org/10.1093/icb/44.6.413.
Boschi, E. E. (2000). Species of decapod crustaceans and their distribution in the American marine zoogeographic provinces. Revista de Investigación y Desarrollo Pesquero, 13, 1–136.
Boschi, E. E. (2002). Distribution of continental shelf decapod crustaceans along the American Pacific Coast. In E. Escobar-Briones & F. Álvarez (Eds.), Modern approaches to the study of Crustacea (pp. 235–239). New York: Springer.
Boyce, D. G., Frank, K. T., & Leggett, W. C. (2015). From mice to elephants: overturning the “one size fits all” paradigm in marine plankton food chains. Ecology Letters, 18(6), 504–515. https://doi.org/10.1111/ele.12434.
Brown, J. H. (1995). Macroecology. Chicago: University of Chicago Press.
Brun, P. G., Payne, M. R., & Kiørboe, T. (2016). Trait biogeography of marine copepods – an analysis across scales. Ecology Letters, 19(12), 1403–1413. https://doi.org/10.1111/ele.12688.
Burnham, K. P., & Anderson, D. R. (2002). Model selection and multimodel inference: a practical information-theoretic approach. New York: Springer.
Cardoso, R., & Defeo, O. (2003). Geographical patterns in reproductive biology of the pan-American sandy beach isopod Excirolana braziliensis. Marine Biology, 143, 573–581. https://doi.org/10.1007/s00227-003-1073-0.
Cepeda, D., Álvarez-Castillo, L., Hermoso-Salazar, M., Sánchez, N., Gómez, S., & Pardos, F. (2019a). Four new species of Kinorhyncha from the Gulf of California, eastern Pacific Ocean. Zoologischer Anzeiger, 282, 140–160. https://doi.org/10.1016/j.jcz.2019.05.011.
Cepeda, D., Sánchez, N., & Pardos, F. (2019b). First report of the family Zelinkaderidae (Kinorhyncha: Cyclorhagida) for the Caribbean Sea, with the description of a new species of Triodontoderes Sørensen & Rho, 2009 and an identification key for the family. Zoologischer Anzeiger, 282, 116–126. https://doi.org/10.1016/j.jcz.2019.05.017.
Cepeda, D., Trigo, D., Pardos, F., & Sánchez, N. (2020). Does sediment composition sort kinorhynch communities? An ecomorphological approach through geometric morphometrics. Scientific Reports, 10, e2603. https://doi.org/10.1038/s41598-020-59511-4.
Chaudhary, C., Saeedi, H., & Costello, M. J. (2016). Bimodality of latitudinal gradients in marine species richness. Trends in Ecology & Evolution, 31, 670–676. https://doi.org/10.1016/j.tree.2016.06.001.
Cheung, W. L., Pauly, D., & Sarmiento, J. L. (2013a). How to make progress in projecting climate change impacts. ICES Journal of Marine Science, 70, 1069–1074. https://doi.org/10.1093/icesjms/fst133.
Cheung, W. L., Sarmiento, J. L., Dunne, J., Frolicher, T. L., Lam, V. W. Y., Deng-Palomares, M. L., Watson, R., & Pauly, D. (2013b). Shrinking of fishes exacerbates impacts of global ocean changes on marine ecosystems. Nature Climate Change, 3, 254–258. https://doi.org/10.1038/nclimate1691.
Chown, S. L., & Gaston, K. J. (2010). Body size variation in insects: a macroecological perspective. Biological Reviews, 85, 139–169. https://doi.org/10.1111/j.1469-185X.2009.00097.x.
Crawley, M. J. (2012). The R book (2nd ed.). Chichester: Wiley.
Crickmore, M. A., & Mann, R. S. (2009). The control of size in animals: insights from selector genes. Bioessays, 30(9), 843–853. https://doi.org/10.1002/bies.20806.
Cushman, J. H., Lawton, J. H., & Manly, B. F. J. (1993). Latitudinal patterns in European ant assemblages: variation in species richness and body size. Oecologia, 95, 30–37. https://doi.org/10.1007/BF00649503.
Defeo, O., & Cardoso, R. S. (2002). Macroecology of population dynamics and life history traits of the mole crab Emerita brasiliensis in Atlantic sandy beaches of South America. Marine Ecology Progress Series, 239, 169–179. https://doi.org/10.3354/meps239169.
Dworschak, P. C. (2000). Global diversity in the Thalassinidea (Decapoda). Journal of Crustacean Biology, 20(2), 238–245. https://doi.org/10.1163/1937240X-90000025.
Escribano, R., Hidalgo, P., Valdés, V., & Frederick, L. (2014). Temperature effects on development and reproduction of copepods in the Humboldt Current: the advantage of rapid growth. Journal of Plankton Research, 36(1), 104–116. https://doi.org/10.1093/plankt/fbt095.
Fenchel, T., & Finlay, B. J. (2004). The ubiquity of small species: patterns of local and global diversity. BioScience, 54(8), 777–784. https://doi.org/10.1641/0006-3568(2004)054[0777:TUOSSP]2.0.CO;2.
Finlay, B. J., & Esteban, G. F. (2007). Body size and biogeography. In A. G. Hildrew, D. G. Raffaelli, & R. Edmonds-Brown (Eds.), Body size: the structure and function of aquatic ecosystems (pp. 167–185). Cambridge: Cambridge University Press.
Foissner, W. (2006). Biogeography and dispersal of microorganisms: a review emphasizing protists. Acta Protozoologica, 45(2), 111–136.
Fontaneto, D., Barbosa, A. M., Segers, H., & Pautasso, M. (2012). The “rotiferologist” effect and other global correlates of species richness in monogonont rotifers. Ecography, 35, 174–182. https://doi.org/10.1111/j.1600-0587.2011.06850.x.
Forster, J., Hirst, A. G., & Atkinson, D. (2012). Warming-induced reductions in body size are greater in aquatic than terrestrial species. PNAS, 109(47), 19310–19314. https://doi.org/10.1073/pnas.1210460109.
Fox, J., & Weisberg, S. (2019). An R companion to applied regression (3rd ed.). Thousand Oaks: Sage.
Freckelton, R. P., Harvey, P. H., & Pagel, M. (2002). Phylogenetic analysis and comparative data: a test and review of the evidence. The American Naturalist, 160, 712–726. https://doi.org/10.1086/343873.
Grzelak, K., & Sørensen, M. V. (2018). Diversity and distribution of Arctic Echinoderes species (Kinorhyncha: Cyclorhagida), with the description of one new species and a redescription of E. arlis Higgins 1966. Marine Biodiversity, 49, 1131–1150. https://doi.org/10.1007/s12526-018-0889-2.
Guillera-Arroita, G., Kéry, M., & Lahoz-Monfort, J. J. (2019). Inferring species richness using multispecies occupancy modelling: estimation performance and interpretation. Ecology and Evolution, 9(2), 780–792. https://doi.org/10.1002/ece3.4821.
Hawkins, B. A. (2011). Eight (and a half) deadly sins of spatial analysis. Journal of Biogeography, 39, 1–9. https://doi.org/10.1111/j.1365-2699.2011.02637.x.
Hillebrand, H., & Azovsky, A. I. (2001). Body size determines the strength of the latitudinal diversity gradient. Ecography, 24, 251–256. https://doi.org/10.1034/j.1600-0587.2001.240302.x.
Higgins, R. P. (1988). Kinorhyncha. In R. P. Higgins & H. Thiel (Eds.), Introduction to the study of meiofauna (pp. 328–331). Washington, DC: Smithsonian Institution Press.
Ho, C. K., Pennings, S. C., & Carefoot, T. H. (2009). Is diet quality an overlooked mechanism for Bergmann’s rule? The American Naturalist, 175, 269–276. https://doi.org/10.1086/649583.
Holt, B., & Jønsson, K.A. (2014) Reconciling hierarchical taxonomy with molecular phylogenies. Systematic Biology, 63, 1010–1017. https://doi.org/10.1093/sysbio/syu061
Horne, C. R., Hirst, A. G., Atkinson, D., Neves, A., & Kiørbe, T. (2016). A global synthesis of seasonal temperature-size responses in copepods. Global Ecology and Biogeography, 25(8), 988–999. https://doi.org/10.1111/geb.12460.
Huston, M. A., & Wolverton, S. (2011). Regulation of animal size by eNPP, Bergmann’s rule and related phenomena. Ecological Monographs, 81, 349–405. https://doi.org/10.1890/10-1523.1.
Kennish, M. J. (2017). Ecology of estuaries, volume II: biological aspects. Boca Ratón: CRC Press.
Kingsolver, J. G., & Huey, R. B. (2008). Size, temperature, and fitness: three rules. Evolutionary Ecology Research, 10(2), 251–268.
Kosnik, M. A., Jablonski, D., Lockwood, R., & Novack-Gottshall, P. M. (2006). Quantifying molluscan body size in evolutionary and ecological analyses: maximizing the return on data-collection efforts. Palaios, 21(6), 588–597. https://doi.org/10.2110/palo.2006.p06-012r.
Kristensen, R. M., & Higgins, R. P. (1991). Kinorhyncha. In R. W. Harrison & E. E. Ruppert (Eds.), Microscopic anatomy of invertebrates, Vol. 4 (pp. 377–404). New York: John Wiley and Sons.
Lindsey, C. C. (1966). Body sizes of poikilotherm vertebrates at different latitudes. Evolution, 20, 456–465.
Lonsdale, D. J., & Levinton, J. S. (1985). Latitudinal differentiation in copepod growth: an adaptation to temperature. Ecology, 66, 1397–1407. https://doi.org/10.2307/1938002.
Margulis, L., & Chapman, M. (2009). Kingdom and domains: an illustrated guide to the phyla of life on earth. Boston: Academic Press.
Martorelli, S., & Higgins, R. P. (2004). Kinorhyncha from the stomach of the shrimp Pleoticus muelleri (Bate, 1888) from Comodoro Rivadavia, Argentina. Zoologischer Anzeiger, 243, 85-98.https://doi.org/10.1016/j.jcz.2004.07.003
Maze, G. (2011). 1997-2007 SeaWIFS based monthly standard estimate of NPP. Available at: http://www.ifremer.fr/lpo/files/gmaze/data/standard_vgpm.seawifs.global.nc.gz (accessed 1 November 2018).
McClain, C. R., & Rex, M. (2001). The relationship between dissolved oxygen concentration and maximum size in deep-sea turrid gastropods: an application of quantile regression. Marine Biology, 139, 681–685. https://doi.org/10.1007/s002270100617.
McDowall, R. M. (2007). Jordan’s and other ecogeographical rules, and the vertebral number in fishes. Journal of Biogeography, 35(3), 501–508. https://doi.org/10.1111/j.1365-2699.2007.01823.x.
McNab, B. K. (2010). Geographic and temporal correlations of mammalian size reconsidered: a resource rule. Oecologia, 164, 13–23. https://doi.org/10.1007/s00442-010-1621-5.
Meyer, K. M., Memiaghe, H., Korte, L., Kenfack, D., Alonso, A., & Bohannan, B. J. M. (2018). Why do microbes exhibit weak biogeographic patterns? The Multidisciplinary Journal of Microbial Ecology, 12(6), 1404–1413. https://doi.org/10.1038/s41396-018-0103-3.
Moran, A. L., & Woods, H. A. (2012). Why might they be giants? Towards an understanding of polar gigantism. The Journal of Experimental Biology, 215, 1995–2002. https://doi.org/10.1242/jeb.067066.
Neira, C., Ingels, J., Mendoza, G., Hernández-López, E., & Levin, L. A. (2018). Distribution of meiofauna in bathyal sediments influenced by the oxygen minimum zone off Costa Rica. Frontiers in Marine Science, 5, e448. https://doi.org/10.3389/fmars.2018.00448.
Neuhaus, B. (2013). Kinorhyncha (= Echinodera). In A. Schmidt-Rhaesa (Ed.), Handbook of Zoology. Gastrotricha, Cycloneuralia and Gnathifera. Volume 1: Nematomorpha, Priapulida, Kinorhyncha, Loricifera (pp. 181–348). Göttingen: De Gruyter.
Neuhaus, B., Pardos, F., Sørensen, M. V., & Higgins, R. P. (2013). Redescription, morphology and biogeography of Centroderes spinosus (Reinhard, 1881) (Kinorhyncha, Cyclorhagida) from Europe. Cahiers de Biologie Marine, 54(1), 109–131. https://doi.org/10.21411/CBM.A.8E3FD0CA.
Pardos, F., Herranz, M., & Sánchez, N. (2016). Two sides of a coin: the phylum Kinorhyncha in Panama (II). Pacific Panama. Zoologischer Anzeiger, 265(26-47), 26–47. https://doi.org/10.1016/j.jcz.2016.06.006.
Partridge, L., & Coyne, L. A. (1997). Bergmann’s rule in ectotherms: is it adaptative? Evolution, 51, 632–635.
Peters, R. H. (1983). The ecological implications of body size. Cambridge: Cambridge University Press.
Pinheiro, J., Bates, D., DebRoy, S., Sarkar, D., & R Core Team (2020). nlme: linear and nonlinear mixed effects models. R package version 3.1–145, https://CRAN.R-project.org/package=nlme. Accessed 1 Feb 2020
R Core Team. (2020). R: a language and environment for statistical computing. Vienna: R Foundation for Statistical Computing http://www.R-project.org/. Accessed 1 Feb 2020
Rex, M. A., Crame, J. A., Stuart, C. T., & Clarke, A. (2005). Large-scale biogeographic patterns in marine mollusk: a confluence of history and productivity? Ecology, 86, 2288–2297. https://doi.org/10.1890/04-1056.
Reynolds, R. W., Rayner, N. A., Smith, T. M., Stokes, D. C., & Wang, W. (2002). An improved in situ and satellite SST analysis for climate. Journal of Climate, 15, 1609–1625. https://doi.org/10.1175/1520-0442(2002)015<609:AIISAS>2.0.CO;2.
Rollinson, N., & Locke, R. (2018). Temperature-dependent oxygen limitation and the rise of Bergmann’s rule in species with aquatic respiration. Evolution, 2018, 977–988. https://doi.org/10.1111/evo.13458.
Rodríguez, A., Olalla-Tárraga, M. A., & Hawkins, B. A. (2008). Bergmann’s rule and the geography of mammal body size in the Western Hemisphere. Global Ecology and Biogeography, 17, 274–283. https://doi.org/10.1111/j.1466-8238.2007.00363.x.
Rosenzweig, M. L. (1968). The strategy of body size in mammalian carnivores. The American Midland Naturalist, 80(2), 299–315. https://doi.org/10.2307/2423529.
Roy, K. (2002). Bathymetry and body size in marine gastropods: a shallow water perspective. Marine Ecology Progress Series, 237, 143–149. https://doi.org/10.3354/meps237143.
Roy, K., Jablonski, D., & Martien, K. K. (2000). Invariant size-frequency distributions along a latitudinal gradient in marine bivalves. PNAS, 97, 13150–13155. https://doi.org/10.1073/pnas.97.24.13150.
Roy, K., Jablonski, D., Valentine, J. W., & Rosenberg, G. (1998). Marine latitudinal diversity gradients: tests of casual hypotheses. PNAS, 95(7), 3699–3702. https://doi.org/10.1073/pnas.95.7.3699.
Saeedi, H., Costello, M. J., Warren, D., & Brandt, A. (2019). Latitudinal and bathymetrical species richness patterns in the NW Pacific and adjacent Arctic Ocean. Scientific Reports, 9, e9303. https://doi.org/10.1038/s41598-019-45813-9.
Saeedi, H., Dennis, T. E., & Costello, M. J. (2017). Bimodal latitudinal species richness and high endemicity of razor clams (Mollusca). Journal of Biogeography, 44, 592–604. https://doi.org/10.1111/jbi.12903.
San Martín, E., Harris, R. P., & Irigoien, X. (2006). Latitudinal variation in plankton size spectra in the Atlantic Ocean. Deep Sea Research Part II: Topical Studies in Oceanography, 53, 1560–1572. https://doi.org/10.1016/j.dsr2.2006.05.006.
Sánchez, N., Herranz, M., Benito, J., & Pardos, F. (2012). Kinorhyncha from the Iberian Peninsula: new data from the first intensive sampling campaigns. Zootaxa, 3402, 24–44. https://doi.org/10.11646/zootaxa.3402.1.2.
Sánchez, N., Pardos, F., Herranz, M., & Benito, J. (2011). Pycnophies dolichurus sp. nov. and P. aulacodes sp. nov. (Kinorhyncha, Honalorhagida, Pycnophyidae), two new kinorhynchs from Spain with a reevaluation of homalorhagid taxonomic characters. Helgoland Marine Research, 65, 319–334. https://doi.org/10.1007/s10152-010-0226-z.
Sánchez, N., Rho, H. S., Min, W. G., Kim, D., & Sørensen, M. V. (2013). Four new species of Pycnophyes (Kinorhyncha: Homalorhagida) from Korea and the East China Sea. Scientia Marina, 77(2), 353–380. https://doi.org/10.3989/scimar.03769.15A.
Sargis, E. J., Millien, V., Woodman, N., & Olson, L. E. (2018). Rule reversal: ecogeographical patterns of body size variation in the common treeshrew (Mammalia, Scandentia). Ecology and Evolution, 8(3), 1634–1645.
Saunders, R. A., & Tarling, G. A. (2018). Southern Ocean mesopelagic fish comply with Bergmann’s rule. The American Naturalist, 191(3), 343–351.
Sibly, R. M., & Atkinson, D. (1994). How rearing temperature affects optimal adult size in ectotherms. Functional Ecology, 8, 486–493. https://doi.org/10.2307/2390073.
Smith, K. F., & Brown, J. H. (2002). Patterns of diversity, depth range and body size among pelagic fishes along a gradient of depth. Global Ecology and Biogeography, 11, 313–322. https://doi.org/10.1046/j.1466-822X.2002.00286.x.
Sørensen, M. V., Heiner, I., & Hansen, J. G. (2008). A comparative morphological study of the kinorhynch genera Antygomonas and Semnoderes (Kinorhyncha: Cyclorhagida). Helgoland Marine Research, 63, 129–147. https://doi.org/10.1007/s10152-008-0132-9.
Sørensen, M. V., Herranz, M., Rho, H. S., Min, W. G., Yamasaki, H., Sánchez, N., & Pardos, F. (2012). On the genus Dracoderes Higgins & Shirayama, 1990 (Kinorhyncha: Cyclorhagida) with a redescription of its type species, D. abei, and a description of a new species from Spain. Marine Biology Research, 8(3), 210–232. https://doi.org/10.1080/17451000.2011.615328.
Sørensen, M. V., & Pardos, F. (2020). Kinorhyncha. In A. Schmidt-Rhaesa (Ed.), Guide to the identification of marine meiofauna (pp. 391–414). Munich: Verlag Dr. Friedrich Pfeil.
Stillwell, R. C. (2010). Are latitudinal clines in body size adaptive? Oikos, 119, 1387–1390. https://doi.org/10.1111/j.1600-0706.2010.18670.x.
Tidière, M., Lemaître, J. F., Pélabon, C., Gimenez, O., & Gaillard, J. M. (2017). Evolutionary allometry reveals a shift in selection pressure on male horn size. Journal of Evolutionary Biology, 30, 1826–1835. https://doi.org/10.1111/jeb.13142.
Tomczak, M. (2019). Upper ocean mean horizontal structure. In J. K. Cochran, H. J. Bokuniewicz, & P. L. Yager (Eds.), Encyclopedia of ocean sciences (3rd edition), volume 1: Marine biogeochemistry (pp. 60–70). Cambridge: Elsevier Academic Press.
Vinarski, M. V. (2014). On the applicability of Bergmann’s rule to ectotherms: the state of the art. Biology Bulletin Reviews, 4, 232–242. https://doi.org/10.1134/S2079086414030098.
Virgós, E., Kowalczyk, R., Trua, A., De Marinis, A., Mangas, J. G., Barea-Azcón, J. M., & Geffen, E. (2011). Body size clines in the European badger and the abundant centre hypothesis. Journal of Biogeography, 38, 1546–1556. https://doi.org/10.1111/j.1365-2699.2011.02512.x.
Walters, R. J., & Hassall, M. (2006). The temperature-size rule in ectotherms: may a general explanation exist after all? The American Naturalist, 167(4), 510–523. https://doi.org/10.1086/501029.
Whittaker, R. J., & Fernández-Palacios, J. M. (2006). Island biogeography; ecology, evolution and conservation (1st ed.). Oxford: Oxford University Press.
Williams, G. C. (1966). Adaptation and natural selection. Princeton: Princeton University Press.
Wyatt, T., & Carlton, J. T. (2002). Phytoplankton introductions in European coastal waters: why are so few invasions reported? CIESM Workshop Monographs, 20, 41–46.
Young, C. M., Arellano, S. M., Hamel, J. F., & Mercier, A. (2006). Chapter 16. Ecology and evolution of larval dispersal in the deep sea. In T. Carrier, A. Reitzel, & A. Heyland (Eds.), Evolutionary ecology of marine invertebrate larvae (pp. 229–250). Oxford: Oxford University Press Scholarship Online.
Zamora-Camacho, F. J., Reguera, S., & Moreno-Rueda, G. (2014). Bergmann’s rules body size in an ectotherm: heat conservation in a lizard along a 2200-metre elevational gradient. Journal of Evolutionary Biology, 27(12), 2820–2828. https://doi.org/10.1111/jeb.12546.
Zuur, A. F., Ieno, E. N., Walker, N. J., & Savelieve, A. A. (2009). Mixed effects models and extensions in ecology with R (1st ed.). New York: Springer.
Funding
DC was supported by a predoctoral fellowship from the University Complutense of Madrid (CT27/16-CT28/16). NS was funded by the Community of Madrid and the University Complutense of Madrid in the framework of the Research Talent Attraction Programme for incorporation into research groups in the Community of Madrid (2019) (2019-T2/AMB-13328).
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FP, NS and DC conceived together the general idea of the study. DC created the dataset and conducted the whole experimental and statistical process. FP, NS and DC discussed the results and defined the main conclusions of the study. DC wrote the manuscript. All the authors reviewed the manuscript.
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Additional Supporting Information may be found in the online version of this article:
ESM 1
Appendix S1 Dataset containing original measures used to extract body size per kinorhynch species, latitude and longitude values defining grids of 1° latitude/longitude, and body size values. Abbreviations: BS, body size; LAT, latitude; LON, longitude; MSW, maximum sternal width; TL, total trunk length. Body measurements are indicated in μm and depth in metres (XLSX 143 kb)
ESM 2
Appendix S2 Supporting tables with the results of the generalized least squares, linear and mixed models analysing size-latitude trends per hemisphere and coastline (PDF 572 kb)
ESM 3
Appendix S3 Supporting tables with the results of the comparisons between generalized least squares models, including five different spatial structures (exponential, Gaussian, spherical, linear and rational quadratic), and the linear models without spatial structure testing the effect of latitude (both raw and squared values), environmental and confounding variables on kinorhynch body size, also including the subdivision into six major coastlines (PDF 511 kb)
ESM 4
Appendix S4 Supporting figures representing graphically size-latitude trends and relationships between body size and environmental variables for the global analyses, and the analyses conducted separately per hemisphere and coastline (only those of statistically significant results are included) (PDF 748 kb)
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Cepeda, D., Pardos, F. & Sánchez, N. From biggest to smallest mud dragons: size-latitude trends in a group of meiobenthic animals worldwide. Org Divers Evol 21, 43–58 (2021). https://doi.org/10.1007/s13127-020-00471-y
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DOI: https://doi.org/10.1007/s13127-020-00471-y