Climate-Driven Range Extension of Amphistegina (Protista, Foraminiferida): Models of Current and Predicted Future Ranges

PLoS ONE

Volumn 8, Issue 2, 2013, Pages

  Langer, Martin R ^a   Weinmann, Anna E ^a   Lötters, Stefan ^b   Bernhard, Joan M ^c   Rödder, Dennis ^d  

^a UNIVERSITY OF BONN   (Germany)

^b UNIVERSITY OF TRIER   (Germany)

^c WOODS HOLE OCEANOGRAPHIC INSTITUTION   (United States)

^d ZOOLOGISCHES FORSCHUNGSMUSEUM ALEXANDER KOENIG   (Germany)

Author keywords

[No Author keywords available]

Indexed keywords

AMPHISTEGINA; AREA UNDER THE CURVE; ARTICLE; CLIMATE CHANGE; CONTROLLED STUDY; ECOLOGICAL NICHE; ENVIRONMENTAL CHANGE; FORAMINIFER; GLOBAL CLIMATE; MEDITERRANEAN SEA; SOUTH AFRICA; SPECIES DISTRIBUTION; SPECIES DIVERSITY; SPECIES EXTINCTION; SPECIES IDENTIFICATION; SYMBIONT; WARMING; WATER TEMPERATURE; AFRICA; BIOLOGICAL MODEL; ECOSYSTEM; FOSSIL; GROWTH, DEVELOPMENT AND AGING;

AFRICA; CLIMATE CHANGE; ECOSYSTEM; FORAMINIFERA; FOSSILS; MEDITERRANEAN SEA; MODELS, BIOLOGICAL;

EID: 84873559572     PISSN: None     EISSN: 19326203     Source Type: Journal    
DOI: 10.1371/journal.pone.0054443     Document Type: Article

References (119)

1. Hoegh-Guldberg O, Bruno JF, (2010) The impact of climate change on the world's marine ecosystems. Science 328: 1523-1528.
2. Tittensor DP, Mora C, Jetz W, Lotze HK, Ricard D, et al. (2010) Global patterns and predictors of marine biodiversity across taxa. Nature 466: 1098-1101.
3. Yamano H, Sugihara K, Nomura K, (2011) Rapid poleward range expansion of tropical reef corals in response to rising sea surface temperatures. Geophys Res Lett 38: L04601 doi:10.1029/2010GL046474.
4. Langer MR, Weinmann AE, Lötters S, Rödder D, (2012) "Strangers" in Paradise: Modeling the biogeographic range expansion of the foraminifera Amphistegina in the Mediterranean Sea. J Foramin Res 42: 234-244.
5. Simberloff D (2000) Introduced species: the threat to biodiversity and what can be done. American Institute of Biological Sciences. Available: http://www.actionbioscience.org/biodiversity/simberloff.html via the Internet. Accessed: 21March 2006.
6. Dale VH, Joyce LA, McNulty S, (2001) Climate change and forest disturbances. BioScience 51: 723-734.
7. Parmesan C, (2006) Ecological and evolutionary responses to recent climate change. Annu Rev Ecol Evol Syst 37: 637-669.
8. LeRoux PC, McGeoch MA, (2008) Rapid range expansion and community reorganization in response to warming. Glob Chang Biol 14: 2950-2962.
9. Root TL, Price JT, Hall KR, Schneider SH, Rosenzweig C, et al. (2003) Fingerprints of global warming on wild animals and plants. Nature 421: 57-60.
10. Parmesan C, Yohe G, (2003) A globally coherent fingerprint of climate change impacts across natural system. Nature 421: 37-42.
11. Clark RM, Thompson R, (2010) Predicting the impact of global warming on the timing of spring flowering. Int J Climatol 30: 1599-1613.
12. Cudmore TJ, Björklund N, Carroll AL, Lindgren BS, (2010) Climate change and range expansion of an aggressive bark beetle: evidence of higher beetle reproduction in nayi{cyrillic}ve host tree populations. J Appl Ecol 47: 1036-1043.
13. Sorte CJB, Williams SL, Carlton JT, (2010) Marine range shifts and species introductions: comparative spread rates and community impacts. Glob Ecol Biogeograph 19: 303-316.
14. Langer MR, Hottinger L, (2000) Biogeography of selected "larger" foraminifera Micropaleontology 46 supplement no. 1: 105-126.
15. Murray JW (1973) Distribution and ecology of living benthic foraminiferids. London: Heinemann Educational Books. p. 244.
16. Zmiri A, Kahan D, Hochstein S, Reiss Z, (1974) Phototaxis and thermotaxis in some species of Amphistegina (Foraminifera). J Protozool 21: 133-138.
17. Larsen AR, (1976) Studies of recent Amphistegina. Taxonomy and some ecological aspects. Israel J Earth Sci 25: 1-26.
18. Langer MR (2008) Foraminifera from the Mediterranean and the Red Sea. In: Por FD, editor. Aqaba-Eilat, the Improbable Gulf: Environment, Biodiversity and Preservation. Jerusalem: Magnes Press. 399-417.
19. Lee JJ, (1995) Living sands. BioScience 45: 252-261.
20. Hallock P, (1981) Production of carbonate sediments by selected large benthic foraminifera on two Pacific coral reefs. J Sediment Petrol 51: 467-474.
21. Langer MR, Silk MT, Lipps JH, (1997) Global ocean carbonate and carbon dioxide production: the role of reef foraminifera. J Foramin Res 27: 271-277.
22. Langer MR, (2008) Assessing the contribution of foraminiferal protists to global ocean carbonate production. J Eukaryot Microbiol 55: 163-169.
23. Hallock P, (1981b) Light dependence in Amphistegina. J Foramin Res 11: 40-46.
24. Yamano H, Miyajima T, Koike I, (2000) Importance of foraminifera for the formation and maintenance of a coral sand cay. Coral Reefs 19: 51-58.
25. Seidel DJ, Fu Q, Randel WJ, Reichler TJ, (2008) Widening of the tropical belt in a changing climate. Nat Geosci 1: 21-24.
26. Fu Q, Johanson CM, Wallace JM, Reichler T, (2006) Enhanced mid-latitutde tropospheric warming in satellite measurements. Science 312: 1179.
27. Pandolfi JM, Connoly SR, Marshall DJ, Cohen AL, (2011) Projecting coral reef futures under global warming and ocean acidification. Science 333: 418-422.
28. Triantaphyllou MV, Dimiza MD, Koukousioura O, Hallock P, (2012) Observations on the life cycle of the symbiont-bearing foraminifer Amphistegina lobifera Larsen, an invasive species in coastal ecosystems of the Aegean Sea (Greece, E. Mediterranean). J Foramin Res 42: 143-150.
29. Triantaphyllou MV, Koukousioura O, Dimiza MD, (2009) The presence of Indo-Pacific symbiont-bearing foraminifer Amphistegina lobifera in Greek coastal ecosystems (Aegean Sea, eastern Mediterranean). Med Mar Sci 10: 73-85.
30. IPCC (2007) Climate change 2007: impacts, adaptation and vulnerability. Contribution of Working Group II to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change. Parry ML, Canziani OF, Palutikof J, van der Linden PJ, Hanson CE, editors. Cambridge and New York: Cambridge University Press. 976 p.
31. Southward AJ, Hawkins SJ, Burrows MT, (1995) Seventy years' observations of changes in distribution and abundance of zooplankton and intertidal organisms in the western English Channel in relation to rising sea temperature. J Therm Biol 20: 127-155.
32. Oviatt CA, (2004) The changing ecology of temperate coastal waters during a warming trend. Estuaries 27: 895-904.
33. Perry AL, Low PJ, Ellis JR, Reynolds JD, (2005) Climate change and distribution shifts in marine fishes. Science 308: 1912-1915.
34. Dambach J, Rödder D, (2011) Applications and future challenges in marine species distribution modelling. Aquat Conserv 21: 92-100.
35. Robinson LM, Elith J, Hobday AJ, Pearson RG, Kendall BE, et al. (2011) Pushing the limits in marine species distribution modelling: lessons from the land present challenges and opportunities. Glob Ecol Biogeograph 20: 789-802.
36. Kintisch E, (2008) Impacts research seen as next climate frontier. Science 322: 182-183.
37. Hannah L, Midgley GF, Millar D, (2002) Climate change integrative conservation strategies. Glob Ecol Biogeograph 11: 485-495.
38. Saikkonen K, Taulavuori K, Hyvönen T, Gundel PE, Hamilton CE, et al. (2012) Climate change-driven species' range shifts filtered by photoperiodism. Nat Clim Change 2: 239-242.
39. Alory G, Wijffels S, Meyers G, (2007) Observed temperature trends in the Indian Ocean over 1960-1999 and associated mechanisms. Geophys Res Lett 34: L02606 doi:10.1029/2006GL028044.
40. Grosholz ED, (1996) Contrasting rates of spread for introduced species in terrestrial and marine systems. Ecology 77: 1680-1686.
41. Kinlan BP, Hastings A (2005) Rates of population spread and geographic expansion. What exotic species tell us. In: Sax DF, Stachowicz JJ, Gaines SD, editors. Species Invasions: Insights into Ecology, Evolution and Biogeography. Sunderland, Massachusetts: Sinauer & Associates. 381-419.
42. Möbius KA (1880) Foraminiferen von Mauritius. In: Möbius KA, Richters F, Martens E, editors. Beiträge zur Meeresfauna der Insel Mauritius und der Seychellen. Berlin: Gutmann. 65-136.
43. Brady HP, (1884) Report on the foraminifera dredged by H.M.S. Challenger, during the years 1873-1876. Reports on the Scientific Results of the Voyage of the H.M.S. Challenger during the years 1873-1876. Zoology 9: 1-814.
44. Egger JG, (1893) Foraminiferen aus Meeresgrundproben, gelotet von 1874 bis 1876 von S.M. Sch. Gazelle. Königliche Bayerische Akademie der Wissenschaften, Mathematisch-Physikalische Klasse, Abhandlungen 18: 139-458.
45. Heron-Allen E, Earland A, (1915) The foraminifera of the Kerimba Archipelago (Portuguese East Africa). Part II. Trans Zool Soc London 20: 543-794.
46. Braga JM, (1961) Foraminíferos da costa de Moçambique. Publicações do Instituto de Zoologia "Dr. Augusto Nobre". Faculdade de ciências do Porto 77: 1-208.
47. Le Calvez Y, (1965) Les foraminifères. In: Cahiers du Office de la Recherche Scientifique et Technique Outre-mer Guilcher A, Berthois L, Le Calvez Y, Battistini R, Crosnier A, editors. Les recifs coralliens et le lagon de l'ile Mayotte (Archipel des Comores, Oceans Indien). 11: 181-201.
48. Moura AR, (1965) Foraminiferos da Ilha da Inhaca. Revista dos Estudos gerais universitarios de Mozambique, Série 2, Ciencias Biológicas e Agranómicas 2: 1-74.
49. Monier C, (1973) Note préliminaire sur les foraminifères benthiques du platier interne du grand récif de Tuléar (Madagascar). Téthys 5: 241-250.
50. Battistini R, Gayet J, Jouannic C, Labracherie M, Peypouquet JP, et al. (1976) Etude des sédiments de la microfaune des Iles Glorieuses (Canal de Mozambique). Cahiers du Office de la Recherche Scientifique et Technique Outre-mer, Série Geologie 8: 147-171.
51. Phleger FB, (1976a) Holocene ecology in St. Lucia lagoon, Zululand based on foraminifera. The St. Lucia scientific advisory council workshop meeting. Charters Creek, 15-17 February 1976: 1-8.
52. Phleger FB, (1976b) Foraminiferal and ecological processes in St. Lucia Lagoon, Zululand. Maritime Sediments, Special Publication 1: 195-204.
53. Hottinger L, (1977) Foraminifères operculiniformes. Mémoires du Muséum National d'Histoire Naturelle, Paris, Série C, Sciences de la terre 40: 1-159.
54. Pereira CPG (1979) Foraminiferal distribution and ecology in the fringing reef complex of the coast, near Mombasa, Kenya. Unpublished PhD thesis, Aberystwyth: University of Wales. 480 p.
55. Chasens SA, (1981) Foraminifera of the Kenya coastline. J Foramin Res 11: 191-202.
56. Montaggioni LF, (1981) Les associations de foraminifères dans les sédiments de récifaux de l'archipel des Mascareignes (Océan Indien). Ann Inst Oceanograph 57: 41-62.
57. Lévy A, Mathieu R, Poignant A, Rosset-Moulinier M, Rouvillois A, (1982) Contribution à la connaissance des foraminifères du littoral Kenyan (Océan Indien). Cah Micropal 2: 135-148.
58. Neagu T, (1982) Foraminifères récents de la zone du recif coralligène de l'île de Mbudya (Côte orientale de la Tansanie). Rev Esp Micropal 14: 99-136.
59. McMillan IK, (1986) Cainozoic planktonic and larger foraminifera distributions around southern Africa and their implications for past changes of oceanic water temperatures. S Afr J Sci 82: 66-69.
60. Cooper JAG, McMillan IK, (1987) Foraminifera of the Mgeni Estuary, Durban and their sedimentological significance. South Afr J Geol 90: 489-498.
61. Wright CI, McMillan IK, Mason TR, (1990) Foraminifera and sedimentation patterns in St. Lucia Estuary mouth, Zululand, South Africa. S Afr J Geol 93: 592-601.
62. Dale DC, McMillan IK (1998) Mud belt and middle shelf benthonic and planktonic foraminiferal assemblages and sedimentation processes compared through the Holocene succession at two tropical African (Sierra Leone) and two temperate African (Western offshore, South Africa) sites. S Afr J Sci, 94, 319-340.
63. Dale DC, McMillan IK (1999) On the Beach: A Field Guide to the Late Cenozoic Micropalaeontological History, Saldanha Region, South Africa. Cape Town: De Beers Marine. 127 p.
64. Perry CT, (2003) Coral reefs in a high-latitude, siliciclastic barrier island setting: reef framework and sediment production at Inhaca Island, southern Mozambique. Coral Reefs 22: 485-497.
65. Perry CT, Beavington-Penney SJ, (2005) Epiphytic calcium carbonate production and facies development within sub-tropical seagrass beds, Inhaca Island, Mozambique. Sediment Geol 174: 161-176.
66. Langer MR, Schmidt-Sinns J, (2006) Biogeography of modern benthic foraminifera from South Africa, Namibia and Mozambique. Ann Inst Geocien 29: 686-687.
67. Pignatti J, Frezza V, Benedetti A, Carbone F, Accordi G, Matteucci R, (2012) Recent foraminiferal assemblages from mixed carbonate-siliciclastic sediments of southern Somalia and eastern Kenya. Boll Soc Geol It 131: 47-65.
68. Hijmans RJ, Guarino L, Rojas E (2001) DIVA-GIS. A geographic information system for the analysis of biodiversity data. Manual: International Potato Center, Lima.
69. Tyberghein L, Verbruggen H, Pauly K, Troupin C, Mineur F, et al. (2011) Bio-ORACLE: a global environmental dataset for marine species distribution modelling. Glob Ecol Biogeograph 21: 272-281.
70. Christensen JH, Hewitson B, Busuioc A, Chen X, Gao I, et al. (2007) Regional climate projections. In: Solomon S, Qin D, Manning M, Chen Z, Marquis M, et al., editors. Climate Change 2007: The Physical Science Basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge and New York: Cambridge University Press. 847-940.
71. Phillips SJ, Dudík M, Shapire RE (2004) A maximum entropy approach to species distribution modeling. Proceedings of the 21st International Conference on Machine Learning: 655-662.
72. Phillips SJ, Anderson RP, Schapire RE, (2006) Maximum entropy modeling of species geographic distributions. Ecol Modell 190: 231-259.
73. Jaynes ET, (1957) Information theory and statistical mechanics. Phys Rev 106: 620-630.
74. Phillips SJ, Dudik M, Elith J, Graham CH, Lehmann A, et al. (2009) Sample selection bias and presence-only distribution models: implications for background and pseudo-absence data. Ecol Appl 19: 181-197.
75. Elith J, Phillips SJ, Hastie T, Dudík M, En Che Y, et al. (2011) A statistical explanation of Maxent for ecologists. Divers Distrib 17: 43-57.
76. Phillips SJ, Dudík M, (2008) Modeling of species distributions with Maxent: new extensions and a comprehensive evaluation. Ecography 31: 161-175.
77. Pearce J, Ferrier S, (2000) An evaluation of alternative algorithms for fitting species distribution models using logistic regression. Ecol Modell 128: 128-147.
78. Ward DF, (2007) Modelling the potential geographic distribution of invasive ant species in New Zealand. Biol Invasions 9: 723-735.
79. Swets JA, (1988) Measuring the accuracy of diagnostic systems. Science 240: 1285-1293.
80. Liu C, Pam M, Dawson TP, Pearson RG, (2005) Selecting thresholds of occurrence in the prediction of species distributions, Ecography. 28: 385-393.
81. d'Orbigny A, (1826) Tableau méthodique de la classe des Céphalopodes. Ann Sci Nat 7: 245-314.
82. Wright EP, (1876) Notes on a small collection of Foraminifera from the Seychelles. Proc R Ir Acad 2: 586-588.
83. Wright EP, (1877) Notes on Foraminifera (from the Seychelles and from Cagliari). J Nat Hist Serie 4 19: 40-44.
84. Banner FT, Pereira C (1981) A temporal and spatial analysis of foraminiferal diversity from the fringing reefs of Mombasa, East Africa. In: Neale JW, Brasier MD, editors. Microfossils from Recent and Fossil Shelf Seas. Chichester and NewYork: Ellis Horwood. 350-366.
85. Albani AD, (1965) The foraminifera in a sample dredged from the vicinity of Salisbury Island, Durban Bay, South Africa. Contrib Cushman Found Foramin Res 16: 60-66.
86. McMillan IK (1974) Recent and relict foraminifera from Agulhas Bank, South African Continental Margin. Unpublished Master thesis, Aberystwyth: University College Wales.
87. Toefy R, Gibbons MJ, McMillan IK, (2005) The foraminifera associated with the alga Gelidium pristoides, South Africa. Afr Invertebr 46: 1-26.
88. Belderson RH (1961) The size distribution characteristics of the Recent shallow marine sediments off Durban, South Africa. Unpublished Master thesis, Durban: University of Natal.
89. IPCC (2007b) Climate change 2007: the physical science basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change. Solomon S, Qin D, Manning M, Chen Z, Marquis M, Averyt KB, Tignor M, Miller HL, editors. Cambridge and New York: Cambridge University Press. 996 p.
90. Belanger CL, Jablonski D, Roy K, Berke SK, Krug AZ, et al. (2012) Global environmental predictors of benthic marine biogeographic structure. Proc Natl Acad Sci U S A 109: 14046-12051.
91. Hiddink JG, (2012) Ben Rais Lasram F, (2012) Cantrill J, (2012) Davies AJ, (2012) Keeping pace with climate change: what can we learn from the spread of Lessepsian migrants? Glob Chang Biol 18: 2161-2172.
92. Caley MJ, Carr MH, Hixon MA, Hughes TP, Jones GP, et al. (1996) Recruitment and the local dynamics of open marine populations. Annu Rev Ecol Evol Syst 27: 477-500.
93. Hallock P, Cottey TL, Forward LB, Halas J, (1986) Population biology and sediment production of Archaias angulatus (Foraminiferida) in Largo Sound, Florida. J Foramin Res 16: 1-8.
94. Goldstein S, (1999) Foraminifera: A Biological Overview. In: Sen Gupta BK, editor. Modern Foraminifera. Dordrecht: Kluwer Academic Publishing. 37-56.
95. Funk C, Dettinger,MD, Michaelsen JC, Verdin JP, Brown ME, et al. (2008) Warming of the Indian Ocean threatens eastern and southern African food security but could be mitigated by agricultural development. Proc Natl Acad Sci U S A 105: 11081-11086.
96. Levitus S, Antonov JI, Boyer TP, Stephens C, (2000) Warming of the world ocean. Science 287: 2225-2229.
97. Barnett TP, Pierce DW, Schnur R, (2001) Detection of anthropogenic climate change in the world's oceans. Science 292: 270-274.
98. Hansen J, Sato M, Ruedy R, Lo K, Lea DW, et al. (2006) Global temperature change. Proc Natl Acad Sci U S A 103: 14288-14293.
99. Cai W, Cowan T, Dix M, Rotstayn L, Ribbe J, et al. (2007) Anthropogenic aerosol forcing and the structure of temperature trends in the southern Indian Ocean. Geophys Res Lett 34: L14611 doi:10.1029/2007GL030380.
100. Rögl F, Brandstätter F, (1993) The foraminifera genus Amphistegina in the Korytnica Clays (Holy Cross Mts., Central Poland) and its significance in the Miocene of the Paratethys. Acta Geol Pol 43: 121-146.
101. Todd R (1976) Some observations about Amphistegina (foraminifera). Progress in Micropaleontology, Selected papers in honor of Prof. Kiyoshi Asano, Micropaleontology Press Special Publication: 382-394.
102. Chaproniere GCH, (1980) Influence of plate tectonics on the distribution of late Paleogene to early Neogene larger foraminiferids in the Australasian region. Palaeogeogr Palaeoclimatol Palaeoecol 31: 299-317.
103. Chaproniere GCH, (1984) Oligocene and Miocene larger Foraminiferida from Australia and New Zealand: Canberra, Commonwealth of Australia, Department of National Resources and Energy, Bureau of Mineral Resources, Geology and Geophysics, Bulletin. 188: 1-98.
104. Scheibner C, Speijer RP, (2008a) Decline of coral reefs during late Paleocene to early Eocene global warming. eEarth 3: 19-26.
105. Scheibner C, Speijer RP, (2008b) Late Paleocene-early Eocene Tethyan carbonate platform evolution- A response to long- and short-term paleoclimatic change. Earth Sci Rev 90: 71-102.
106. Goldbeck E, Langer MR, (2009) Biogeographic provinces and patterns of diversity in selected Upper Cretaceous (Santonian-Maastrichtian) larger foraminifera. In: SEPM Special Publication Demchuk T, Gary AC, editors. Geological Problem Solving with Microfossils: A Volume in Honor of Garry D. Jones. 93: 187-232.
107. Pujalte V, Schmitz B, Baceta JI, Orue- Etxebarria X, Bernaola G, et al. (2009) Correlation of the Thanetian-Ilerdian turnover of larger foraminifera and the Paleocene-Eocene thermal maximum: confirming evidence from the Campo area (Pyrenees, Spain). Geologica Acta 7: 161-175.
108. Zachos JC, Pagani M, Sloan L, Thomas E, Billups K, (2001) Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292: 686-693.
109. Bohaty SM, Zachos JC, (2003) Significant Southern Ocean warming event in the late middle Eocene. Geology 31: 1017-1020.
110. Pearson PN, (2010) Increased atmospheric CO2 during the Middle Eocene. Science 330: 763-764.
111. Hallock P (1999) Symbiont-bearing foraminifera. In: Sen Gupta BK, editor. Modern Foraminifera. Dordrecht: Kluwer Academic Publishing. 123-140.
112. Hallock P, Triantaphyllou MV, Dimiza MD, Koukousioura O, (2011) Invasive foraminifera in coastal ecosystems of the eastern Mediterranean: Implications for understanding larger foraminiferal-dominated biofacies in the Cenozoic. GSA, Abstracts with Programs 43: 556.
113. McIntyre-Wressnig A, Bernhard, JM, McCorkle DC, Hallock P (in press) Non-lethal effects of ocean acidification on the symbiont-bearing benthic foraminifera Amphistegina gibbosa. Mar Ecol Prog Ser. in press.
114. Vogel N, Uthicke S, (2012) Calcification and photobiology in symbiont-bearing benthic foraminifera and responses to a high CO2 environment. J Exp Mar Bio Ecol 424: 15-24.
115. Abu Tair NK (2011) Der Impakt invasiver Foraminiferen aus dem Roten Meer auf das mediterrane Ökosystem von Zypern. Unpublished Diploma thesis, Bonn: Rheinische Friedrich-Wilhelms-Universität Bonn. 73 p.
116. Jones CG, Lawton JH, Shachak M, (1994) Organisms as ecosystem engineers. Oikos 69: 373-386.
117. Jones CG, Lawton JH, Shachak M, (1997) Positive and negative effects of organisms as physical ecosystem engineers. Ecology 78: 1946-1957.
118. Lüning K (1990) Seaweeds: Their Environment, Biogeography and Ecophysiology. New York: Wiley-Interscience. p. 527.
119. Schmidt-Nielsen K (1990) Temperature effects. In: Schmidt-Nielsen K, editor. Animal Physiology: Adaptation and Environment. New York: Cambridge University Press. 217-239.