Placing populations and their attributes into a geographic context is currently the thing to do. Partly this may be because mapping has been revolutionized by geographic information systems (GIS) technology and the increasing power of desktop computers. Also, molecular data now allow inference of monophyla that are worth mapping; and for population-level analyses, there is phylogeography (), a statistically rigorous way of overlaying geography onto an estimated gene tree to measure the strength of geography/phylogeny associations. Age estimation from molecular sequences has emerged as another powerful new tool.

With access to absolute times, evolution can be linked to geological events and, for the first time, the most recent arrival of a lineage in an area can be estimated, which is different from the information gained from fossils. Inferring the historical assembly of ecological communities via the comparison of multiple dated phylogenies is a recent outgrowth of this ability (; ). Lastly, the geographic mosaic theory of coevolution (), although hardly full-fledged (), may have added further to the excitement about the geographic context of evolution and adaptation. This new edition of Biogeography by Mark V.

Lomolino, Brett R. Riddle, and James H. Brown thus comes at an opportune time.

The first edition of Biogeography by Brown and Arthur C. Gibson was published in 1983, the second edition, by Brown and Lomolino, 15 years later in 1998, and the third a mere 6 years later in 2005. This is not just a textbook—it is the most comprehensive text and general reference book in the field, now with a 50-page long bibliography that cites over 1000 sources published between 1820 and 2004. There are 18 chapters, grouped into six units, and 447 black-and-white illustrations, mostly graphs and maps, but also a few wonderful photos of people and landscapes. Unit 1, focusing on the history of biogeography, has hardly changed from the previous editions, and ends with the role of null hypotheses in studies of community assembly. Unit 2 includes chapters on the physical setting (solar radiation, winds, rainfall, soils, aquatic environments, oceans), the factors covering distributions of single species and those that may govern the geography of communities. As expected, there are brief descriptions of all major biomes.

Unit 3, “Earth History and Fundamental Biogeographic Processes,” has a chapter on dispersal (focusing on autecology and with only 19 post-1998 references), and one on speciation and extinction. The latter includes discussions of species concepts, micro- and macroevolution, and modes of speciation. This year's astonishing discovery of a case of sympatric speciation in palms on an oceanic island () unfortunately could not yet be included. In an idiosyncratic choice, Unit 3 has the geological time scale, continental drift, and Pleistocene glaciations (Chapters 8 and 9) following speciation and extinction (Chapter 7).

Many of the paleodistribution maps are new, and the text of all four chapters has been much updated. For teaching purposes, the Northern Hemisphere responses to the Pleistocene climatic cycles are the ideal basis for a discussion of global warming, and the book's bias towards its largest market is nowhere more evident than in this chapter, which contains a single figure illustrating Pleistocene changes outside the Americas (p. Unit 4, “Evolutionary History of Lineages and Biotas,” comprises chapters on endemism and biogeographic regions, on reconstructing the history of lineages and that of biotas.

*Correspondence: Mark V. Lomolino, College of Environmental Science and Forestry. Syracuse, NY 13031, USA. E-mail: island@esf.edu. Modern biogeography now encompasses an impressive diversity of patterns and phenomena of the geography of nature, providing insights fundamental to understanding. Refer the reader to our earlier paper. (Lomolino and Brown 1989) for a more extended description of Munroe's inde- pendent development of an equilibrium theory of island biogeography (see also ex- cerpts from his dissertation available at the. International Biogeography Society web site, http://www.biogeography.org/html/.

The last two are the contribution of the new coauthor of Biogeography, Brett Riddle, whose research focuses on the phylogeography of Great Basin montane island biota and molecular systematics of North American rodents. He has done an excellent job of explaining the relevant basic concepts, such as phylogenetic inference, properties of molecular characters, construction and interpretation of haplotype networks, and molecular clocks. The sections explaining the role of fossils in biogeography are all new, and very good. By comparison with another recently revised text, Cox and Moore's Biogeography (cf. The review by ), Lomolino et al. Provide much more information about methods, such as reconciling trees, dispersal-vicariance analysis and Brooks parsimony analysis, always pointing the reader to appropriate original literature.

Unit 5 turns to ecological biogeography, and comprises two chapters on island biogeography—Brown and Lomolino are important contributors to the current nonequilibrium view of island biota—and a chapter on diversity gradients and macroecology from the keyboard of Brown. This chapter pays tribute to historical explanations of diversity, even if the exposition is strangely a historical. For example, the tropical conservatism hypothesis (that the tropics on average are larger and older than other biomes and regions, and that speciation rates there are higher and extinction rates lower so that species over time accumulated in the tropics), which is beautifully set out in Herbert Baker's (1970) classic review “Evolution in the Tropics” and also detailed by and other authors in the Ricklefs and Schluter book on species diversity, is attributed to a 2004 paper in Trends in Ecology and Evolution (). Those people who are interested in explanations of diversity disparities up to 1980, of course, can turn to Foundations of Biogeography: Classic Papers with Commentaries, a fascinating compendium. Foundations and Frontiers () have been produced by the young and fit International Biogeography Society, whose cofounder and first president was Lomolino. (Brown and Riddle also are cofounders and past and present presidents of the 6-year-old society.) The final Unit (6), “Conservation Biogeography and New Frontiers,” comprises chapters on the biodiversity crisis, invasive species, conservation biogeography, and the biogeography of humans.

The book ends with a chapter on the frontiers of biogeography, excerpted from Frontiers. The only omission in this book seems to be a section on GIS methods of mapping and model-based analyses of species ranges. GIS is mentioned in the Introduction and on the very last pages (p.

722, 747; these mentions are not in the index), but the capabilities and use of widely available packages, such as Domain, BIOCLIM, and GARP, and their role in Element Distribution Modeling, is something that the next generation of biogeographers, to be taught with this book, really needs to know about. Nowhere in the index does one find predictive distribution modeling, predictive range mapping, species distribution mapping, habitat distribution mapping, or ecological niche modeling. The authors of Biogeography hold that little in evolution, and for that matter ecology, paleontology, conservation biology, and human evolution, makes sense unless viewed in a geographic context. The new edition of their book strongly supports this contention, and the very personal, sometimes chatty, sometimes missionary style in which it is written conveys a feeling that one is close to people who have seen the action.

This is an empirically and conceptually rich text (as Andy Sinauer stated about the second edition), whose third edition confirms its status as an indispensable classic.

• Department of Environmental and Forest Biology, SUNY College of Environmental Science and Forestry, Syracuse, New York, USA E-mail: • BIOSKETCH Mark V. Lomolino's research and teaching focus on biogeography and conservation of biological diversity. He received the American Society of Mammalogist Award, and serves on the editorial advisory board for Biological Conservation, is an Editor of Global Ecology and Biogeography: A Journal of Macroecology, and is coauthor with James H. Brown of Biogeography (published by Sinauer Associates). • • `there are areas too small, and areas too large, to show clear diversity patterns': 191) Comment on ) The species–area relationship does not have an asymptote. Journal of Biogeography, 28, 827–830. Inertia is a fundamental property of nature and, in a very real sense, an important force in the development of science.

Even during periods of conceptual stasis, inertial resistance to paradigmatic shifts temper the development of a discipline, resisting careless adoption of just any new and novel theory that challenges the champion. Eventually, however, our understanding of the complexity of nature stretches far beyond the elastic limits of the long-standing paradigm. At such times, inertial resistance to alternative and unconventional views may impede the advance of scientific disciplines, sometimes for decades.

The case of such delayed scientific development that I am most familiar with is that of the equilibrium theory of island biogeography. Eugene Gordon Munroe's early articulation of the equilibrium theory in 1948 was ignored by his colleagues, yet it was conceptually equivalent to that proposed and finally accepted as the paradigm of the field of island biogeography in the late 1960s (see,;,; ). Central to the acceptance of the equilibrium theory of island biogeography was the paradigmatic shift from the view of island biotas as static assemblages of species, to one where insular communities are viewed as dynamic in time—varying in their species composition due to recurrent immigrations and extinctions. It was likely more than just coincidence that it was also in the late 1960s that scientists finally accepted the theory of continental drift and plate tectonics, which holds that the earth itself is dynamic. The theory was first formerly proposed by Alfred Wegener and F. Taylor in the early 1900s (; ), but was resisted and often ridiculed by most `respectable' scientists until the accumulated burden of information on the dynamics of earth and its biotas finally snapped the long-standing doctrine of geological stasis; but not until some five decades after Taylor and Wegener published their challenges to the static theory, and centuries after ) first commented on the fit of the continents. These lessons in delayed development of scientific theories characterizes the dynamic tension between accepted, theoretical constructs and our ever-expanding appreciation for the complexity of nature.

The topic of ) critique of my article () on the `protean' and possible sigmoidal nature of the species–area relationship may not be as monumental as the paradigm shifts discussed above, but some of the same lessons may apply. In the 1920s, ) and ) proposed alternative mathematical formulae to describe and analyse the species–area relationship—both depicting curves with positive, albeit attenuating slopes. In 1962, Preston canonized Arrhenius' log–log, or `power' model and its exponent ` z' as the formula of choice for most biogeographers.

Yet, in his long and rich double-manuscript on the species–area relationship (,), he discussed an alternative form of the species–area relationship—the sigmoidal curve (1962a: 187, 214–215). Later, in their seminal monograph on island biogeography,: 32) referred to the peculiarities of biotas on small islands such as the plants of Kapingamarangi Atoll, Polynesia () and ants of Micronesia (). For such small islands, species-richness seemed to vary independently of island area—a pattern inconsistent with the conventional mathematical models (power or semi-log) of the species–area relationship. These conventional models also do not allow for the possible contribution of in situ (within island) speciation to insular biotas which, as Munroe suggested in 1953, should contribute to a secondary increase in the slope of the species–area relationship on very large islands. As Robert MacArthur concluded in his final monograph, Geographical Ecology (: 191), `The general conclusion of this section [on environmental scale] is that there are areas too small, and areas too large, to show clear diversity patterns, but that for the proper intermediate census area, the patterns are clear.'

Thus, my suggestion that the species–area relationship may be sigmoidal (; fig. 1), and therefore fundamentally different from the models most of us have been using for decades, may still appear heretical, but not as original as it may have first seemed. Again quoting Preston (1962: 215), `the possibility that such [sigmoidal] curves may exist can hardly be disputed on theoretical grounds; how often they occur in practice is a matter for observation.' Recently, ), and ) have provided some very relevant `observations' on the potential sigmoidal nature of the species–area relationship—adding their voices to the epistemological rumblings (see also ). Michael Weiser and I have followed up my challenges to the existing paradigm (, ) with a review and analysis of the form of the species–area relationship in just over 100 insular biotas. Sigmoidal species–area relationships appear to be relatively common, and it seems that alternative analytical models can be used to investigate fundamentally different components of the species–area relationship (see ).

) raise some interesting questions, but their view appears to be based largely on visual inspection of three graphs and their assertions that (i) the species–area relationship does not have an asymptotic or limited richness, and (ii) that there is no small island effect. First, once the species pool is defined, the maximum number of species can not exceed this value, whether we call it an asymptote or boundary. Perhaps this is just a semantic difference, and one that appears much more subtle than my challenge to assess the importance of the small island effect, the potential sigmoidal nature of the species–area relationship, and a possible secondary increase in the species–area curve where in situ speciation may become relevant (: fig. 1; see also ).

That is, it appears that the key distinction between conventional models and the newer alternatives (,;; ) is that the latter include scale-dependent phase shifts. Along gradients of increasing area, as different structuring forces come into play and take prominence, the nature of the species–area curve changes. The `thresholds', or island sizes at which these phase shifts occur, have biological and practical relevance and therefore should receive increasing attention from ecologists and biogeographers. These and other responses to my calls for alternative models to investigate this very fundamental pattern, and criticisms of related views presented in the above-mentioned papers, are to be expected and embraced because they form the fabric of the filters that control and direct the development of our science. In this particular case, however, I believe it is time to modify the traditional, paradigmatic models – the power and semi-log models – in favour of a model which includes a means to test for potential, sigmoidal patterns and phase shifts. As discussed in recent papers (, ), the primary advantage of these alternative models is that they provide an objective means to study the scale-dependent forces structuring insular communities—what: 186) termed the `hierarchical structure' of the `real environment.'

These scale-dependent forces include hurricanes and other disturbances that may strongly influence community structure especially on the smaller islands, to more deterministic, ecological factors and processes associated with habitat diversity, carrying capacity and immigration/extinction dynamics on islands of intermediate size, and finally to speciation on islands large enough to provide the geographical isolation (within islands) necessary for evolutionary divergence. As we have shown (), `alternative' models for the species–area relationship can include these thresholds while retaining basic elements of the conventional models (semi-log and power models). Thus, in addition to studying z-values and other parameters of the traditional models, the modified models can shed light on some important questions.

How should the first threshold (i.e. The range of the small island effect) and the second threshold (marking islands where speciation becomes a prominent structuring force) vary among biotas or among types of archipelagoes? What proportion of islands fall within the range of the small island effect, where area is a poor predictor of diversity, and how much does richness vary within this range? For all its early heuristic value and long-standing tenure as the reigning, paradigmatic model, the power model provides no means to examine such questions. To the extent to which we believe these are important questions, we should continue to search for alternative approaches for studying one of ecology's most fundamental patterns (see also;; ).

If these and other challenges to the existing paradigm prove fruitless, so be it, but continued adherence to a traditional paradigm should be based on repeated challenges and re-evaluation, and not on inertia and acceptance by default. ACKNOWLEDGMENTS Michael D. Weiser provided useful comments on an earlier version of this manuscript. I thank the members of the Foundations of Biogeography Working Group, National Center for Ecological Analysis and Synthesis (USA) for their stimulating and provocative discussion of some of the ideas presented here. Footnotes • BIOSKETCH Mark V.

Lomolino's research and teaching focus on biogeography and conservation of biological diversity. He received the American Society of Mammalogist Award, and serves on the editorial advisory board for Biological Conservation, is an Editor of Global Ecology and Biogeography: A Journal of Macroecology, and is coauthor with James H. Brown of Biogeography (published by Sinauer Associates).

Ancillary Article Information. • 1 Arrhenius, O. ( 1921) Species and area. Journal of Ecology, 4, 68– 73. • • 2 Brown, J.H.

& Lomolino, M.V. ( 1989) On the nature of scientific revolutions: independent discovery of the equilibrium theory of island biogeography. Ecology, 70, 1954– 1957. • • • 3 Brown, J.H. & Lomolino, M.V. ( 2000) Concluding remarks: historical perspective and the future of island biogeography theory.

Global Ecology and Biogeography, 9, 87– 92. • • • 4 Crawley, M.J. & Harral, J.E.

( 2001) Scale dependence in plant biodiversity. Science, 291, 864– 868. • • • • • 5 Gleason, H.A. ( 1922) On the relation between species and area.

Ecology, 3, 158– 162. • • • 6 Heaney, L.R. ( 2000) Dynamic disequilibrium: along-term, large-scale perspective on the equilibrium model of island biogeography.

Global Ecology and Biogeography, 9, 59– 74. • • • 7 Lomolino, M.V. ( 2000a) Ecology's most general, yet protean pattern: the species–area relationship. Journal of Biogeography, 27, 17– 26.DOI: • • • 8 Lomolino, M.V. ( 2000b) A call for a new paradigm of island biogeography. Global Ecology and Biogeography, 9, 1– 6.DOI: • • • 9 Lomolino, M.V.

( 2000c) A species-based theory of insular zoogeography. Global Ecology and Biogeography, 9, 39– 58.DOI: • • • 10 Lomolino, M.V.

( 2001) The species–area relationship: new challenges for an old pattern. Progress in Physical Biogeography, 25, 1– 21. • • 11 Lomolino, M.V. & Weiser, M.D. ( 2001) Towards a more general species–area relationship: diversity on all islands, great and small. Journal of Biogeography, 28, 431– 445.DOI: • • • 12 Losos, J.B.

& Schluter, D. ( 2000) Analysis of an evolutionary species–area relationship.

Nature, 408, 847– 850. • • • • • 13 MacArthur, R.H. ( 1972) Geographical ecology: patterns in the distributions of species. Harper & Row, New York, NY.

• 14 MacArthur, R.H. & Wilson, E.O. ( 1963) An equilibrium theory of insular zoogeography. Evolution, 17, 373– 387.

• • • 15 MacArthur, R.H. & Wilson, E.O. ( 1967) The theory of island biogeography.

Monographs in Population Biology, No. Princeton University Press, Princeton, NJ. • 16 Meadows, M. Dark Souls 2 Product Key Free. E. ( 2001) Biogeography: does theory meet practice? Progress in Physical Geography, 25, 134– 142. • • • 17 Munroe, E.G. ( 1948) The geographical distribution of butterflies in the West Indies.

PhD Dissertation, Cornell University, Ithaca, NY. • 18 Munroe, E.G. ( 1953) The size of island faunas. Proceedings of the Seventh Pacific Science Congress of the Pacific Science Association.

IV, Zoology, 52–53. Whitcome and Tombs, Auckland, New Zealand. • 19 Niering, W.A.

( 1963) Terrestrial ecology of Kapingamarangi Atoll, Caroline Islands. Ecological Monographs, 33, 131– 160. • • • 20 Ortelii (Ortelius), A., ( 1596) The compleat geographer: or, The chorography and topography of all the known parts of the earth. (1772 English translation by Herman Moll).

Printed for J. Knapton, London. • 21 Preston, F.W. ( 1962a) The canonical distribution of commonness and rarity: part I. Ecology, 43, 185– 215. • • • 22 Preston, F.W.

( 1962b) The canonical distribution of commonness and rarity: part II. Ecology, 43, 410– 432. • • • 23 Taylor, F.B.

( 1910) Bearing of the Tertiary mountain belt on the origin of the earth's plan. Geography Society of America Bulletin, 21, 179– 226. • • 24 Wegener, A. ( 1912) Die Entstehung der Kontinente. Petermanns Geographyr.

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& Lonsdale, W.M. ( 2001) The species–area relationship does not have an asymptote. Journal of Biogeography, 28, 827– 830.DOI: • • • 27 Wilson, E.O. & Taylor, R.W. ( 1967) An estimate of the potential evolutionary increase in species density in the Polynesian ant fauna. Evolution, 21, 1– 10. • • Related content.