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Biogeographical Challenges to Neo-Darwinian Evolution

Casey Luskin

Defenders of evolution commonly make the assertion that biogeography provides unequivocal support for neo-Darwinian theory. For example, the pro-Darwin National Center for Science Education (NCSE) claims that "consistency between biogeographic and evolutionary patterns provides important evidence about the continuity of the processes driving the evolution and diversification of all life," and "[t]his continuity is what would be expected of a pattern of common descent."

The reality, however, is that the NCSE has dramatically overstated its case.

In making its case for common descent, the NCSE essentially ignores the numerous and significant examples where the biogeographical evidence does not fit well with purported "evolutionary patterns." In other words, they cherry pick the data and ignore examples where there are great discontinuities between biogeography and neo-Darwinism. One of the most significant discontinuities, the origin of South American monkeys ("called platyrrhines"). The NCSE made a specific argument for common descent based upon the "continuity" and "consistency" between biogeography and evolution. The evidence presented below refutes their assertion.

In its response regarding marsupials, the NCSE admits, "If the [North American] opossum truly had roots in Australia, it would indeed be a biogeographic conundrum." Since North American opossums are not descended from Australian "possums," their high morphological similarity dictates to neo-Darwinian evolutionists that this must be another case of extreme convergent evolution that challenges the methodology by which neo-Darwinism infers homology and common descent.

But what if North American opossums were descended from Australian possums? Why does the NCSE observe that this hypothetical situation would pose a "biogeographic conundrum?" The NCSE says this because there would be no route by which Australian possums could have migrated to North America. The NCSE's reasoning here is sound: they presume that if organisms in Locale B are descended from organisms in Locale A, then there must have been some migration route by which organisms could migrate from A to B. If there is no such route, then we're presented with, in the NCSE's own words, a "biogeographic conundrum." Using such reasoning, the NCSE then argues that marsupials and other groups have biogeographic histories that are congruent with the tectonic history of islands and continents, thus allegedly supporting common descent: The same pattern of diversification and migration seen in marsupials can also be seen in other groups of plants and animals. That consistency between biogeographic and evolutionary patterns provides important evidence about the continuity of the processes driving the evolution and diversification of all life. This continuity is what would be expected of a pattern of common descent, and is not what would be expected with the creationist orchard scheme. With marsupials, the NCSE claims that the "continuity" of geography and evolution predicts that there will always be some land bridge or migratory pathway which terrestrial organisms can follow. This was claimed to allegedly show "consistency between biogeographic and evolutionary patterns" that demonstrates "what would be expected of a pattern of common descent." Ignoring the NCSE's continued inappropriate usage of the "creationist" label, their claim is simply not true, for there are many examples of terrestrial organisms existing and appearing in locations where no land-based migratory route is apparent. The NCSE's approach is to cherrypick examples to support their arguments for universal common descent, but a large number of "biogeographic conundrums" that challenge neo-Darwinism could be discussed.

Traditional evolutionary explanations of biogeography fail when terrestrial (or freshwater) organisms appear on an island or continent but there is no standard migratory mechanism for them to have arrived there from some ancestral population. The NCSE boasts about the use of migration pathways or land bridges to explain the presence of marsupials or other plants and animals around the world. But what happens when organisms--even higher mammals--appear on isolated islands, and there appears no way for their purported ancestors to migrate there? At these points, evolutionary biogeographers appeal to a fallback position, a suite of mechanisms of "oceanic dispersal." As a review by De Quieroz (2005) stated: A classic problem in biogeography is to explain why particular terrestrial and freshwater taxa have geographical distributions that are broken up by oceans. Why are southern beeches (Nothofagus spp.) found in Australia, New Zealand, New Guinea and southern South America? Why are there iguanas on the Fiji Islands, whereas all their close relatives are in the New World?57 According to De Quiroz, such examples require "oceanic dispersal over tectonic vicariance as an explanation for disjunct distributions in a wide variety of taxa, from frogs to beetles to baobab trees."58 But he recognizes a fundamental problem with overseas dispersal hypotheses: "cladistic biogeographers claimed that hypotheses of dispersal were not falsifiable because all patterns of relationships can be explained by some dispersal hypothesis."59 He further states that, "A main objection to dispersal hypotheses is that they are unfalsifiable and thus unscientific," continuing that, "this can be countered by noting that, if plausible vicariance hypotheses are falsified, then dispersal is supported by default."60 In other words, evolutionists assume that traditional land-based migration pathways (the type of evidence the NCSE claims supports common descent) were taken, but when they aren't an option, one can always fall back when necessary on to unfalsifiable ad hoc hypotheses of oceanic dispersal. After reviewing a number of "unexpected" biogeographic data that require oceanic dispersal, De Quiroz's review concludes: "these cases reinforce a general message of the great evolutionist [Darwin]: given enough time, many things that seem unlikely can happen."

Thus, neo-Darwinian evolutionists are forced to appeal to "unlikely" or "unexpected" transmigration of terrestrial organisms, in some cases requiring the crossing of oceans ("oceanic dispersal") to account for some biogeographical data. Such data challenges the simplistic picture of biogeography put forth by the NCSE that biogeography lends support to universal common descent through congruence between migration pathways and tectonic history. If anything, the "disjunct distributions in a wide variety of taxa" would tend to lend prima facie support for an orchard model of life's history; a single tree of life hypothesis can only be sustained through extremely unlikely ad hoc appeals to oceanic dispersal to save universal common descent from difficult biogeographical data. What follows are some notable examples of such data.

Sea Monkey Hypotheses
One of the most infamous examples of the very sort of "biogeographic conundrum" the NCSE fears is the origin of South American monkeys, called platyrrhines.61 Based upon molecular and morphological evidence, "New World" platyrrhine monkeys are thought to be descended from African "Old World" or catarrhine monkeys. The problem is that plate tectonic history shows that Africa and South America split off from one another between 100 and 120 Mya, and that South America was an isolated island continent at least from about 80 Mya until about 3.5 Mya.62 Molecular studies claim that the South American monkeys split from African monkeys perhaps around 35 Mya.63 Monkeys are thought to have first evolved in Africa, and so somehow proponents of neo-Darwinism must account for the subsequent appearance of monkeys in the Upper Oligocene in South America.64 As Walter Carl Hartwig puts it: "The platyrrhine origins issue incorporates several different questions. How did platyrrhines get to South America?"65

If the standard evolutionary story is true, and platyrrhines and catarrhines are both part of the same crown group radiation of monkeys, then how did platyrrhines come to be in South America if South America was then an isolated island continent and there was no land-based route for monkeys to migrate from Africa to South America? For those unfamiliar with the arguments that proponents of neo-Darwinian biogeography make when backed into a corner, the answer to these questions is almost too incredible to believe: they propose that monkeys floated on rafts across the Atlantic Ocean to colonize South America. And of course, we can't have just one seafaring monkey, or the monkey will quickly die leaving no offspring. Thus, at least two monkeys (or perhaps a single pregnant monkey) must have made the rafting voyage.

If this proposal seems a little farfetched, consider the quite serious endorsement of the rafting hypothesis given in a recent authoritative book, Primate Biogeography: Progress and Prospects (2006). The authors of the chapter "The Biogeography of Primate Evolution," John G. Fleagle and Christopher C. Gilbert, state the problem as follows: The most biogeographically challenging aspect of platyrrhine evolution concerns the origin of the entire clade. South America was an island continent throughout most of the Tertiary, and most of the orders of mammals found in Paleocene through Miocene deposits are endemic families or orders almost exclusively restricted to that continent. Primates first appear in the Late Oligocene and become common only in the Early Miocene. Rodents also appear first in the Oligocene. Both groups are almost certainly immigrants from some other continent, and paleontologists have debated for much of this century how and where primates reached South America.66 Likewise, a Harper Collins textbook on human evolution states: The origin of platyrrhine monkeys puzzled paleontologists for decades. ... When and how did the monkeys get to South America? Prior to about 1970, paleontologists invoked the concept of parallel evolution. ... It seemed so unlikely that monkeys from Africa could cross a water barrier like the Atlantic Ocean... Molecular evidence demonstrated that all monkeys shared a common ancestor prior to their separation. ... The "rafting hypothesis" argues that monkeys evolved from prosimians once and only once in Africa, and that it is a primitive monkey (parapithecid), and not a prosimian, that made the water-logged trip to South America. ... Other species colonizing South America must have arrived in similar ways over millions of years.67 As noted above, the high degree of molecular genetic similarity between platyrrhine and catarhine monkeys precludes the possibility that African and South American monkeys are similar simply because of convergent evolution. Yet as Fleagle and Gilbert state, similarities between monkeys across the oceans "raises a difficult biogeographical issue" because "South America is separated from Africa by a distance of at least 2600 km, making a phylogenetic and biogeographic link between the primate faunas of the two continents seem very unlikely."68 They argue that in light of "[t]he absence of any anthropoids from North America, combined with the considerable morphological evidence of a South American-African connection with the rodent and primate faunas" that therefore "the rafting hypothesis is the most likely scenario for the biogeographic origin of platyrrines."69

All kinds of arguments have gone back and forth about whether such a rafting journey is possible or plausible. Of course, millions of years ago Africa and South America were slightly closer than they are today, but they were still very far apart at the time monkeys supposedly colonized South America. Fleagle and Gilbert argue that at best, the position of the continents in the early Tertiary still requires a "journey from Africa to South America anywhere from 8 to 15 days."70 This is called "plausible," but a macroview must be taken here: Is there any real biogeographical evidence that can falsify common ancestry? If the presence of higher mammalian fauna on isolated island continents with no simple way to arrive there does not falsify neo-Darwinian explanations of biogeography, what will?

Indeed, the rafting hypothesis has serious problems, for monkeys and rodents have high metabolisms and require large amounts of food and water: The case of platyrrhines is more difficult to explain as anthropoid primates have higher metabolic rates and do not have the ability for prolonged periods of topor. A two-week rafting event across the Atlantic must have involved a floating island with an adequate food and water supply.71 Such "floating islands" are said to exist, but they admit that "the prevalence of over-water dispersal during primate evolution seems truly amazing for a mammalian order."72 They further admit that "[t]he reasons for the prevalence of rafting during the course of primate evolution remain to be explained."73

Needless to say, not all feel comfortable believing that seafaring monkeys on rafts are "plausible." As Hartwig puts it, "The overwhelming evidence for the late Cretaceous-Pliocene isolation of South America renders the mechanical aspect of platyrrhine dispersal virtually irresolvable,"74 for "any late Eocene origins model must invoke a transoceanic crossing mechanism that is implausible (rafting) or suspect (waif dispersal) at best."75

And there are deeper problems: monkeys apparently made the journey, but other smaller African primates such as lorises and galagos never colonized South America. If it was so easy for monkeys to raft across the proto-Atlantic ocean, why didn't these lower primates also make the voyage? The answer we're given by Fleagle and Gilbert is that rafting is "clearly a chance event, an example of 'sweepstakes' dispersal" as "[o]ne can only speculate that by a stroke of good luck anthropoids where able to 'win' the sweepstakes while lorises and galagos did not."76 As another authority wrote, "The evidence strongly suggests the existence of a Palaeogene transoceanic sweepstakes route between Africa and South America, and presumably also a similar route between Africa and Madagascar" to explain such primate distributions.77

Apparently the NCSE was not quite accurate when they claimed, "By comparing macroevolutionary patterns between different groups, we find that the same patterns repeat. This strongly suggests that the same forces drove the diversification of those different groups." The truth is that whenever oceanic "sweepstakes" dispersal is required, we find an exception to expected neo-Darwinian rules of biogeography. And as will be seen in my next post, there are so many exceptions that one might reasonably question whether the inviolable neo-Darwinian rule of universal common ancestry is supported by biogeography.

Sea Monkeys Are the Tip of the Iceberg: More Biogeographical Conundrums for Neo-Darwinism When proponents of neo-Darwinism "speculate" about the "luck" and "chance" needed to explain this "amazing" phenomenon and "challenging" biegeographical data, it's clear that they are lacking reasonable explanations. Yet rafting or other means of "oceanic dispersal" have been suggested to solve a number of other biogeographical conundrums that challenge neo-Darwinism, including:
  • Lizards reaching South America78
  • Large caviomorph rodents reaching South America79
  • Bees arriving in Madagascar80
  • Lemurs arriving in Madagascar81
  • The arrival of other mammals in Madagascar, including the Tenrecidae (hedgehoglike insectivorous mammals), aardvarks, the hippopotamus, and the Viverridae (cat-sized carnivorous mammals)82
  • Dispersal of salamanders across the western end of the Mediterranean83
  • Dispersal of certain lizards across the western end of the Mediterranean84
  • The origin of certain lizards in Cuba85
  • The appearance of elephant fossils on "many islands," which are said to have arrived by swimming86
  • Dispersal of freshwater frogs across oceanic island chains87
  • Certain frogs reaching Madagascar88
  • The colonization of Anguilla by green iguanas89
  • Appearance of certain South American insects90
  • Dispersals of chameleons across the Indian Ocean91
  • Origin of certain insects in Caribbean islands92
  • The origin of mantellid frogs found on the island of Mayotte in the Comoros archipelago, despite the fact that "[a]mphibians are thought to be unable to disperse over ocean barriers because they do not tolerate the osmotic stress of salt water"93
  • The spread of flightless insects to the Chatham Islands94
  • The origin of indigenous gekkos in South America95
  • Origin of crocodile distributions96
  • The appearance of sloths in South America97
  • The origin of a group of Australian rodents98
  • The appearance of land mammals of the Mediterranean islands (also suggesting that "Hippos, elephants, and giant deer reached the islands by swimming")99
  • The origin of various land reptiles in Western Samoa100
  • The presence of Crotalus rattlesnakes in Baja California101
  • Indeed, a review in 2005 by Alan de Queiroz wrote that "[s]triking examples of oceanic dispersal" include: (a) Scaevola (Angiospermae: Goodeniaceae) three times from Australia to Hawaii; (b) Lepidium mustards (Angiospermae: Brassicaceae) from North America and Africa to Australia; (c) Myosotis forget-me-nots (Angiospermae: Boraginaceae) from Eurasia to New Zealand and from New Zealand to South America; (d) Tarentola geckos from Africa to Cuba; (e) Maschalocephalus (Angiospermae: Rapateaceae) from South America to Africa; (f) monkeys (Platyrrhini) from Africa to South America; (g) melastomes (Angiospermae: Melastomataceae) from South America to Africa; (h) cotton (Angiospermae: Malvaceae: Gossypium) from Africa to South America; (i) chameleons three times from Madagascar to Africa; (j) several frog genera to and from Madagascar; (k) Acridocarpus (Angiospermae: Malpighiaceae) from Madagascar to New Caledonia; (l) Baobab trees (Angiospermae: Bombacaceae: Adansonia) between Africa and Australia; (m) 200 plant species between Tasmania and New Zealand; (n) many plant taxa between Australia and New Zealand; and (o) Nemuaron (Angiospermae: Atherospermataceae) from Australia (or Antarctica) to New Caledonia.102 Figure 1 of De Queiroz's paper contains a revealing map of the world covered in lines criss-crossing back and forth across oceans showing how many species must have traversed oceans to explain their distributions in locations unexpected by traditional biogeography:

    deQueiroz2005_fig1.jpg

    (Reprinted from Trends in Ecology and Evolution, Vol.20(2), Alan de Queiroz, "The resurrection of oceanic dispersal in historical biogeography," pages 68-73, (February 2005) with permission from Elsevier. Slightly resized to fit website formatting.)

    It seems clear that there are plenty of examples that contradict the NCSE's simplistic picture of biogeography where the alleged "consistency between biogeographic and evolutionary patterns provides important evidence about the continuity ... [that] would be expected of a pattern of common descent." Somehow all of the above examples got left off the NCSE's commentary on biogeography. There seem to be many more "biogeographic conundrums" than the NCSE is letting on.

    Testing the Orchard Model and the NCSE's Claims of "Nested Patterns" Supporting a "Tree of Life"

    As we have seen, there are many examples where there is inconsistency between evolutionary expectations of biogeography and the established history of plate tectonics. The NCSE is thus wrong to have claimed that "The consistency of these sorts of nested patterns cannot be explained without reference to common descent. The creationist 'orchard' is scientifically meaningless, since it makes no predictions." * The classical "universal common descent" view is contrasted with the orchard model at below:
    treevorchard.jpg
    The NCSE's claim is perplexing because, as noted, the NCSE also claimed that "continuity [between biogeographic and evolutionary patterns] is what would be expected of a pattern of common descent, and is not what would be expected with the creationist orchard scheme." (emphasis added) Ignoring the NCSE's inappropriate use of the "creationist" label, the NCSE seems to be suggesting that the orchard model is contradicted by the data. The NCSE is thus committing the classic evolutionist fallacy of arguing that opposing views are both unfalsifiable, and falsified by the data.

    Regardless, the NCSE is wrong when it claims that the orchard model makes no predictions. If a monophyletic view of common descent predicts "nested patterns," then by the NCSE's own admission, a polyphyletic or "orchard" view predicts non-nested patterns. Indeed, systematists regularly search for precisely such non-nested patterns in order to identify polyphyletic taxa, a phenomenon effectively predicted by the orchard model. The only idea here that is "meaningless" is the NCSE's claim that universal common descent makes predictions, while the "orchard" model does not (and, by the way, is falsified due to its failed predictions).

    Biogeography is full of incongruent patterns which essentially entail non-nested distribution of species. In fact, Bruce S. Lieberman's treatise, Paleobiogeography: Using Fossils to Study Global Change, Plate Tectonics, and Evolution, compares the problem of finding incongruent (i.e., non-nested) patterns among different biogeographic hypotheses to the problem of finding incongruent (i.e., non-nested) patterns of traits in different species when constructing phylogenetic trees: [H]istorical biogeography is the discipline that looks at how groups of organisms have evolved and how their geographic distributions have changed in relation to geological or climatic events. ... In phylogenetic analysis, the arbiter among competing hypotheses suggested by different character systems, i.e. incongruence among characters, is parsimony. The analogous problem in biogeography is what to do when one group suggests one biogeographic pattern, and another group suggests another.103 Analagous to maximizing parsimony in tree-construction, Lieberman notes that vicariance, or land-based separation of organisms, is the preferred explanation, such that the model that can "maximize vicariance, is the one preferred."104 But Lieberman notes that there can be "incongruence"105 between biogeographic patterns. In Lieberman's words, when "one group suggests one biogeographic pattern, and another group suggests another," we have contradictory biogeographical data and we find the opposite of the NCSE's claimed "continuity" that supports universal common descent.

    In this regard, much of the data discussed in my previous post (and the "seamonkey post") entails such incongruence and a breakdown in nested patterns of biogeographic distribution of taxa. As seen, such disparate data often require evolutionists to resort to speculative and unfalsifiable hypotheses of oceanic dispersal as a means of transcending traditional methods of migration. This data challenges the "continuity" of biogeographic and evolutionary patterns said to support universal common descent, but it might be expected under an orchard model.

    In fact, it is not only within biogeography that we find non-nested patterns, and it important to fact-check the NCSE's claim that we always find "nested patterns" pointing to a "tree of life." There's a January 2009 article in New Scientist titled, "Why Darwin Was Wrong about the Tree of Life." Contrary to the NCSE's claim that we always find "nested patterns" which "cannot be explained without reference to common descent," the article reported a major "problem" encountered by molecular systematists, namely that "different genes told contradictory evolutionary stories." The article observed that with the sequencing of the genes and proteins of various living organisms, the tree of life fell apart: "For a long time the holy grail was to build a tree of life," says Eric Bapteste, an evolutionary biologist at the Pierre and Marie Curie University in Paris, France. A few years ago it looked as though the grail was within reach. But today the project lies in tatters, torn to pieces by an onslaught of negative evidence. Many biologists now argue that the tree concept is obsolete and needs to be discarded. "We have no evidence at all that the tree of life is a reality," says Bapteste. That bombshell has even persuaded some that our fundamental view of biology needs to change.106 Of course, these scientists are all committed evolutionists, which makes their admissions all the more weighty. And these arguments apply not just to the base of the tree of life, but also to higher branches where processes like horizontal gene transfer are not thought to be prevalent, as the article observed that "the evolution of animals and plants isn't exactly tree-like either."

    To reiterate, the basic problem is that one gene or protein yields one version of the "tree of life," while another gene or protein yields an entirely different tree. As the New Scientist article stated: The problems began in the early 1990s when it became possible to sequence actual bacterial and archaeal genes rather than just RNA. Everybody expected these DNA sequences to confirm the RNA tree, and sometimes they did but, crucially, sometimes they did not. RNA, for example, might suggest that species A was more closely related to species B than species C, but a tree made from DNA would suggest the reverse.107 Likewise, leading evolutionary bioinformatics specialist W. Ford Doolittle explains, "Molecular phylogenists will have failed to find the 'true tree,' not because their methods are inadequate or because they have chosen the wrong genes, but because the history of life cannot properly be represented as a tree."108 The NCSE may claim that this problem is only encountered when one tries to reconstruct the evolutionary relationships of microorganisms, such as bacteria, which can swap genes through a process called "horizontal gene transfer," thereby muddying any phylogenetic signal. But the tree of life is challenged even among higher organisms where such promiscuous gene-swapping across taxa is not thought to not take place. As the article explains: Syvanen recently compared 2,000 genes that are common to humans, frogs, sea squirts, sea urchins, fruit flies and nematodes. In theory, he should have been able to use the gene sequences to construct an evolutionary tree showing the relationships between the six animals. He failed. The problem was that different genes told contradictory evolutionary stories. This was especially true of sea-squirt genes. Conventionally, sea squirts--also known as tunicates--are lumped together with frogs, humans and other vertebrates in the phylum Chordata, but the genes were sending mixed signals. Some genes did indeed cluster within the chordates, but others indicated that tunicates should be placed with sea urchins, which aren't chordates. "Roughly 50 per cent of its genes have one evolutionary history and 50 per cent another," Syvanen says.109 Even among higher organisms, "[t]he problem was that different genes told contradictory evolutionary stories," leading Syvanen to say, regarding the relationships of these higher groups, "We've just annihilated the tree of life." This directly contradicts the NCSE's claim that there exists a "tree of life" with "nested patterns" which "cannot be explained without reference to common descent."

    Other scientists agree with the conclusions of the New Scientist article. While many try to explain phylogenetic conflicts as being the result of gene swapping among microorganisms at the base of the tree of life, problems with the tree of life extend all the way up the tree. Likewise, Carl Woese, a pioneer of evolutionary molecular systematics, observed that these problems extend well beyond the base of the tree of life: "Phylogenetic incongruities [conflicts] can be seen everywhere in the universal tree, from its root to the major branchings within and among the various taxa to the makeup of the primary groupings themselves."111

    National Academy of Sciences biologist Lynn Margulis has had harsh words for the field of molecular systematics, which Hillis studies. In her article, "The Phylogenetic Tree Topples," she explains that "many biologists claim they know for sure that random mutation (purposeless chance) is the source of inherited variation that generates new species of life and that life evolved in a single-common-trunk, dichotomously branching-phylogenetic-tree pattern!" But she dissents from that view and attacks the evolutionary systematists, noting, "Especially dogmatic are those molecular modelers of the 'tree of life' who, ignorant of alternative topologies (such as webs), don't study ancestors."112

    Striking admissions of troubles in reconstructing the "tree of life" also came from a paper in the journal PLoS Biology entitled, "Bushes in the Tree of Life." The authors acknowledge that "a large fraction of single genes produce phylogenies of poor quality," observing that one study "omitted 35% of single genes from their data matrix, because those genes produced phylogenies at odds with conventional wisdom."113 The paper suggests that "certain critical parts of the [tree of life] may be difficult to resolve, regardless of the quantity of conventional data available."114 The paper even contends that "[t]he recurring discovery of persistently unresolved clades (bushes) should force a re-evaluation of several widely held assumptions of molecular systematics."115

    Unfortunately, one assumption that these evolutionary biologists are not willing to consider changing is the assumption that neo-Darwinism and universal common ancestry are correct. Meanwhile, as far as the data are concerned, the New Scientist article admits, "Ever since Darwin the tree has been the unifying principle for understanding the history of life on Earth," but because "different genes told contradictory evolutionary stories," the notion of a tree of life is now quickly becoming a vision of the past -- as the article stated, it's being "annihilated."

    The NCSE claims that the "orchard" concept is "meaningless" but it would seem to predict the precise non-nested phylogenetic data reported in New Scientist and the non-nested biogeographic data reported in Part III above. Perhaps the reason why different genes are telling "different evolutionary stories" and "one group suggests one biogeographic pattern, and another group suggests another" is because the genes and organisms have wholly different stories to tell, namely stories that indicate that not all living organisms are ancestrally related, thereby fulfilling a testable prediction of the orchard model.

    References Cited:
    * All quotes of the NCSE in this document were downloaded from the NCSE website's response to Explore Evolution on Biogeography on October 29, 2008.

    [57.] Alan de Queiroz, "The resurrection of oceanic dispersal in historical biogeography," Trends in Ecology and Evolution, Vol.20(2):68-73 (February 2005).

    [58.] Id.

    [59.] Id.

    [60.] Id.

    [61.] See John C. Briggs, Global Biogeography, pg. 124 (Elsevier Science, 1995); Alain Houle, "The Origin of Platyrrhines: An Evaluation of the Antarctic Scenario and the Floating Island Model," American Journal of Physical Anthropology, Vol. 109:541--559 (1999).

    [62.] Carlos G. Schrago and Claudia A. M. Russo, "Timing the origin of New World monkeys," Molecular Biology and Evolution, Vol. 20(10):1620--1625 (2003); John J. Flynn and André R. Wyss, "Recent advances in South American mammalian paleontology," Trends in Ecology and Evolution, Vol. 13(11):449-454 (November, 1998); C. Barry Cox & Peter D. Moore, Biogeography: An Ecological and Evolutionary Approach, pg. 185 (Blackwell Science, 1993).

    [63.] Carlos G. Schrago and Claudia A. M. Russo, "Timing the origin of New World monkeys," Molecular Biology and Evolution, Vol. 20(10):1620--1625 (2003).

    [64.] Anthony Hallam, An Outline of Phanerozoic Biogeography, pg. 166 (Oxford University Press, 1994). See also Walter Carl Hartwig, "Patterns, Puzzles and Perspectives on Platyrrhine Origins," in Integrative Paths to the Past: Paleoanthropological Advances in Honor of F. Clark Howell, pg. 80 (Edited by Robert S. Corruccini and Russell L. Ciochon, Prentice Hall, 1994).

    [65.] Walter Carl Hartwig, "Patterns, Puzzles and Perspectives on Platyrrhine Origins," in Integrative Paths to the Past: Paleoanthropological Advances in Honor of F. Clark Howell, pg. 69 (Edited by Robert S. Corruccini and Russell L. Ciochon, Prentice Hall, 1994).

    [66.] John G. Fleagle and Christopher C. Gilbert, "The Biogeography of Primate Evolution: The Role of Plate Tectonics, Climate and Chance," in Primate Biogeography: Progress and Prospects, pgs. 393-394 (Shawn M. Lehman and John G. Fleagle, eds., Springer, 2006) (emphasis added).

    [67.] Adrienne L. Zihlman, The Human Evolution Coloring Book, 4-11 (Harper Collins, 2000).

    [68.] John G. Fleagle and Christopher C. Gilbert, "The Biogeography of Primate Evolution: The Role of Plate Tectonics, Climate and Chance," in Primate Biogeography: Progress and Prospects, pg. 394 (Shawn M. Lehman and John G. Fleagle, eds., Springer, 2006).

    [69.] Id. at 394-395 (emphasis added).

    [70.] Id. at 394.

    [71.] Id. at 404.

    [72.] Id. at 404 (emphasis added).

    [73.] Id. at 403.

    [74.] Walter Carl Hartwig, "Patterns, Puzzles and Perspectives on Platyrrhine Origins," in Integrative Paths to the Past: Paleoanthropological Advances in Honor of F. Clark Howell, pg. 76 (Edited by Robert S. Corruccini and Russell L. Ciochon, Prentice Hall, 1994).

    [75.] Id. at 84 (emphasis added). Note: "waif dispersal" in this case refers to "island-hopping."

    [76.] John G. Fleagle and Christopher C. Gilbert, "The Biogeography of Primate Evolution: The Role of Plate Tectonics, Climate and Chance," in Primate Biogeography: Progress and Prospects, pg. 395 (Shawn M. Lehman and John G. Fleagle, eds., Springer, 2006) (emphases added).

    [77.] Anthony Hallam, An Outline of Phanerozoic Biogeography, pg. 166 (Oxford University Press, 1994).



    [78.] John C. Briggs, Global Biogeography, pg. 93 (Elsevier Science, 1995).

    [79.] Id. at 124.

    [80.] Susan Fuller, Michael Schwarz, and Simon Tierney, "Phylogenetics of the allodapine bee genus Braunsapis: historical biogeography and long-range dispersal over water," Journal of Biogeography, Vol. 32:2135--2144 (2005).

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