The Monkey's Voyage

by Alan de Queiroz

  231The case, as it turned out, was very clear: McDowall (2005).

  231Recent DNA analyses using museum specimens: Austin et al. (2013). For Darwin’s idea of the wolf reaching the islands on icebergs, Darwin (1859).

  233“The Falkland Isld. flora seems to combine . . . ”: Darwin letter to J. D. Hooker, November 28, 1843, from the Darwin Correspondence Project website, www .darwinproject.ac.uk/home. For Croizat linking the Falklands biota to Patagonia, McDowall (2005).

  235A species’ chance of going extinct: For a general review of the effect of area on extinction, Rosenzweig (2002).

  237Graphs plotting the number of extinctions: A good recent study of origination and extinction rates for taxa in the marine fossil record for the past 500 million years is Alroy (2008).

  237“The Great Dying”: Gould (1977). The figure of more than 90 percent extinction for the end-Permian event is in Jablonski (1994).

  238“A severe sorting of the Falklands biota”: McDowall (2005), 59.

  238The ranges of many species . . . shifted to the south: Grayson (1993).

  239The endemic lineages consist of just one or a few species: McDowall (2005).

  239The Chathams . . . are a small archipelago: For the appearance and human history of the Chathams, Harding et al. (2002).

  240Formed part of Zealandia: For the Chathams as part of Zealandia and Gondwana, Campbell and Hutching (2007).

  240The fossil remains of a typical Gondwanan flora: For Late Cretaceous plants and dinosaurs on the Chathams, Stillwell et al. (2006); Campbell and Hutching (2007).

  240The Chathams were completely submerged: For the Cenozoic geological history of the Chathams, including submergence, Campbell et al. (1988); Campbell and Hutching (2007); Stillwell and Consoli (2012).

  241Endemic birds of the Chathams: For the similarity of Chatham Islands and New Zealand birds, Robertson and Heather (2005). A study showing the genetic similarity of the Black Robin of the Chathams to related New Zealand species is Miller and Lambert (2006).

  241Confirmed by extensive molecular studies: Molecular studies of Chatham Islands species are reviewed in Paterson et al. (2006); Goldberg et al. (2008); and Heenan et al. (2010). An apparent anomaly is Myosotidium hortensium, a Chathams plant in the Boraginaceae family, which is estimated to have split from its closest relatives 3.6 to 22.4 million years ago (Goldberg et al. 2008). However, even if the older age limit is correct, this example does not refute the submergence hypothesis, since the divergence did not necessarily occur on the Chathams (see the discussion of New Caledonia later in this chapter). Also, the investigators in this case suggest that this potentially old divergence could be the result of researchers not yet having sampled the closest relatives outside of the Chathams, which seems especially probable, since the nearest known relative is from the Mediterranean region.

  242Amborella trichopoda . . . and the Kagu: For the evolutionary positions of Amborella and the kagu, respectively, Soltis et al. (2008); Fain and Houde (2004). For an entertaining description of the flora and fauna of New Caledonia, see Flannery (1994).

  242New Caledonia was entirely underwater: For geological evidence for the submergence of New Caledonia, Pelletier (2006); Grandcolas et al. (2008).

  242Many New Caledonian lineages, including the skinks: Lygosomine skinks are estimated to have diverged from relatives elsewhere 12.7 to 40.7 million years ago (Smith et al. 2007), which slightly overlaps with the drowning period, but Smith et al. argue that the younger limit is probably more accurate. Araucaria diverged from relatives elsewhere 10.8 to 38.2 million years ago, which also slightly overlaps with the drowning period (Pillon 2012). Other taxa with estimated divergence ages that fall after the drowning include many other plants; Angustonicus cockroaches (formerly considered an ancient relict group); galaxiid fishes; and Paratya freshwater shrimp, among others (Pillon 2012; Grandcolas et al. 2008).

  242Others, such as Amborella: Divergence ages for earliest branching points within ancient New Caledonian groups include the following: diplodactylid geckos, 9 to 21.6 million years (Nielsen et al. 2011); troglosironid harvestmen, 28 to 49 million years (Boyer et al. 2007), 49 million years (Sharma and Giribet 2009), 52 to 102 million years (Giribet et al. 2010), and 40 to 73 million years (Giribet et al. 2012). Amborella trichopoda and the kagu are single species and therefore are unlikely to show ancient divergences within New Caledonia.

  244Similarly unique, ancient lineages: Lactoris, Gamerro and Barreda (2008); Hillebrandia, Clement et al. (2004); Bolyeriidae, Pyron and Burbrink (2011).

  244Compilation of molecular dating studies: Wallis and Trewick (2009).

  244Rails . . . have dispersed: Steadman (1995).

  245Pure vicariance scenario for ratites: For the branching order in the ratite tree, Harshman et al. (2008); Phillips et al. (2010). For molecular dating studies of ratites, Baker and Pereira (2009), which gives estimates from seven studies, and Phillips et al. (2010).

  246The common ancestor of all ratites could fly: Harshman et al. (2008); Phillips et al. (2010). For loss of flight in eighteen living families of birds, Harshman et al. (2008).

  246Two lineages of mite harvestmen: Giribet et al. (2012).

  246Two lineages of centipedes: Murienne et al. (2010).

  246Ironically, New Zealand: For a discussion of the origins of New Zealand’s terrestrial invertebrate fauna, see Giribet and Boyer (2010).

  247We come to Madagascar: For description of Madagascar’s biota, Yoder and Nowak (2006); Goodman and Benstead (2004).

  247When Madagascar detached from India: Krause (2003).

  247Most biologists did not interpret the biota: For origins of Madagascar’s biota by long-distance dispersal, Briggs (1987); Schatz (1996). Others did tend to interpret the biota as a vicariant, relict one. See, for instance, Wickens (1982); Gillespie and Roderick (2002).

  248A key literature-review study: Yoder and Nowak (2006).

  248Colonization and ocean currents: Ali and Huber (2010); Samonds et al. (2012).

  249The key studies have been headed by David Krause: Krause (2003); Krause et al. (2006). The 2003 paper gives the argument for the lack of a connection between the Cretaceous and modern vertebrate faunas.

  250Madagascar and the other Gondwanan islands: In this tour of Gondwanan islands, I have left out Sri Lanka, New Guinea, and Socotra, because those islands have been connected to continents relatively recently.

  250Swarms of igneous dikes: Storey et al. (1999).

  251An archipelago I haven’t discussed: For specific examples in which molecular dating supports overwater colonization of the Seychelles, Austin et al. (2004) on day-geckos; Daniels (2011) on freshwater crabs; and Guo et al. (2012) on wolf snakes.

  251“New Caledonia must be considered . . . ”: Grandcolas et al. (2008), 3309.

  251“Extinction, colonization and speciation . . . ”: Goldberg et al. (2008), 3319.

  252“For me, . . . the particularly interesting aspect of these patterns . . . ”: McDowall (2005), 59.

  253In September 1995, hurricanes Luis and Marilyn: On the story of the green iguanas arrival by rafting, Censky et al. (1998). The status of the iguanas in 2011 is from an email from Ellen Censky to the author, January 10, 2011.

  Chapter Eleven: The Structure of Biogeographic “Revolutions”

  257Some of the caterpillars are amphibious: For amphibious caterpillars and snail-­eating caterpillars, Rubinoff and Schmitz (2010) and Rubinoff and Haines (2005), respectively.

  260Unusually large pseudogenes: Baldo et al. (2011).

  261Their presence in Hawaii: For the distribution of bristletails in Hawaii, Sturm (1993).

  261Quite a few volcanic islands: For the distributions of bristletails in general, including that of Neomachilellus, Sturm and Machida (2001).

  262A 201
2 compilation: Gillespie et al. (2012).

  262Examined the anatomical traits: Sturm (1993). The DNA sequences from Hawaiian specimens are from unpublished work by Robert Meredith, John Gatesy, Cheryl Hayashi, Eric Stiner, and myself.

  262Eggs attached to driftwood: Sturm (1993). For the observation that some bristletail eggs are resistant to chemicals, Larink (1972), cited in Sturm and Bach de Roca (1988).

  263Islands-as-dead-ends rule: Arguments against island-to-mainland colonizations are in Bellemain and Ricklefs (2008).

  263Natural invasions recorded in recent times: Levin (2006).

  264Group includes close to six hundred: O’Grady et al. (2010).

  265Hawaiian escapees have given rise: Published DNA-based phylogenetic evidence for at least one Hawaii-to-mainland dispersal by Scaptomyza, and speculation about why these flies are such good dispersers are in O’Grady and DeSalle (2008). For evidence for other out-of-Hawaii dispersals by Scaptomyza, email from O’Grady to the author, May 23, 2012.

  265Only one of several: For Rhantus beetles, Balke et al. (2009); for Anolis lizards, Nicholson et al. (2005), Glor et al. (2005); for monarchid flycatchers, Filardi and Moyle (2005).

  266A diehard land-bridge advocate: Bowler (1996); Schuchert (1932). The latter includes Schuchert’s argument about granitic rocks on Atlantic islands.

  267Jots down some calculations: Simpson (1952) gives actual calculations of this sort.

  268“Beset by attempts to make the facts fit the theory . . . ”: Email from Briggs to the author, August 25, 2010.

  268Existence of granite on Tristan da Cunha . . . and Ascension: The granitic rock on Ascension Island has been dated as at most a few million years old (Kar et al. 1998), so it clearly has nothing to do with Mesozoic land bridges.

  268Land connections to . . . Hawaii: Skottsberg (1925).

  270The final product, the general area cladogram: For the view that the goal of historical biogeography is to infer relationships among areas, Ebach and Humphries (2002); Ebach (2003); Parenti and Ebach (2009).

  271“Early on I took the vicariance model as the default . . . ”: Email from Trewick to the author, December 13, 2010.

  272It was hard to explain the fact: For anomalies leading to the Copernican revolution, Kuhn (1970).

  273“Regularly marked by frequent and deep debates”: Kuhn (1970), 47–48.

  274“Throughout the pre-paradigm period . . . ”: Kuhn (1970), 163.

  274Matthew’s criticism of land-bridge advocates: Matthew (1915).

  274“Apart from Croizat . . . ”: Email from Heads to the author, August 6, 2010.

  274–275“I don’t think he made . . . ” and “Cladistics by itself . . . ”: Email from Briggs to the author, August 25, 2010.

  275If this view is correct: Mayr (1982, 857), while not dismissing the notion of a scientific revolution, wrote, “I cannot think of a single case in biology where there was a drastic replacement of paradigms between two periods of “normal science.” Mayr was suggesting that Kuhn’s views do not apply well to biology, but his observation can be taken in a different way. Specifically, it may be that some of the great “revolutions” in biology represent the initial emergence of a paradigm that has yet to be replaced, rather than shifts from one accepted paradigm to another. The development of genetics, of cladistic/phylogenetic thinking, and, most conspicuously, of Darwinian evolutionary thinking in general, might be seen in that way.

  276The missing element was time: The importance of placing ages on evolutionary branching points has been emphasized by Donoghue and Moore (2003); de Queiroz (2005); Renner (2005); and Yoder and Nowak (2006), among many others.

  277The great age of many Hawaiian lineages: Skottsberg (1941).

  277“Impervious to evidence”: Yoder, telephone conversation with the author, March 10, 2009.

  277“They were seen for what they are . . . ”: Email from Trewick to the author, December 13, 2010.

  277“I thought, man, I’m having some really weird flashback . . . ”: Donoghue, telephone conversation with the author, September 3, 2010.

  277“Artefactual”: Nelson and Ladiges (2001), 389.

  277“Reactionary”: Santos (2007), 1471.

  277“Ignoring basic biogeographic realities”: McCarthy (2005), 3.

  279On or around October 12, 1988: The description of the locust invasion is in Richardson and Nemeth (1991). The DNA studies indicating that African locusts colonized the New World are Lovejoy et al. (2006); Song et al. (2013).

  Chapter Twelve: A World Shaped by Miracles

  283Let us consider the potato: For the initial cultivation and early South American history of potatoes, Spooner et al. (2005); Mann (2011); McNeill (1999).

  283Ships coming from the Pacific side: McNeill (1999).

  283Rumors dogged the plant: Nunn and Qian (2011).

  283Spread all across northern Europe: McNeill (1999); Nunn and Qian (2011).

  283In Ireland, the impetus: Mann (2011); Pollan (2001).

  283This was the potato blight: Mann (2011); Reader (2008, 2009).

  284The blight reached its peak: Kennedy et al. (2000); Pollan (2001); Mann (2011).

  284Some 2 million people out of Ireland: Mann (2011); Reader (2008).

  284Russia, Germany: McNeill (1999).

  284Rise in population: Nunn and Qian (2011). The population of 600 million in 1700 is from Nunn and Qian, but the figure of 1.5 billion for 1900 subtracts the population of the New World (roughly 100 million) from their figure of 1.6 billion.

  285“Homogenocene”: Charles Mann’s 2011 book 1493 provides an engaging and detailed look at many of the consequences of the intentional and inadvertent introduction of species all over the world since the discovery of the Americas by Europeans. The term “Homogenocene” was coined by Samways (1999).

  285Unremarkable day in the life: This account draws from the following: nutmeg, Joseph (1980); eggplant, Olmstead and Palmer (1997); zucchini and watermelon, Schaefer et al. (2009); common bean, Lavin et al. (2004); corn, Bouchenak-Khelladi et al. (2010); crocodiles, Meredith et al. (2011a); monkeys, Fleagle (1999) and Poux et al. (2006); guinea pigs, Poux et al. (2006) and Rowe et al. (2010); lovebirds, Schweizer et al. (2010); chameleons, Tolley et al. (2013); cotton, Wendell et al. (2010).

  287Ancestor of anthropoids: Chaimanee et al. (2012). All placements of the fossils within the primate evolutionary tree indicate a crossing of the Tethys Sea, but whether the crossing was to or from Africa is unclear.

  288South America was an island continent: McLoughlin (2001); Iturralde-Vinent (2006); Brown et al. (2006); Mann et al. (2007).

  290“Great American Interchange”: Webb (2006). The history of South American mammals through the Cenozoic (the past 66 million years), including the Great American Interchange, was described most famously by George Gaylord Simpson in his 1980 book, Splendid Isolation, but the basic story of evolution in isolation, followed by mixing, had been suggested much earlier, particularly by W. B. Scott in 1932.

  290Island-hopping journeys: For the earliest procyonid fossils in South America, Webb (2006). For the earliest sigmodontine fossils in South America, Verzi and Montalvo (2008). The exact age of the sigmodontine fossils has been questioned by Prevosti and Pardiñas (2009), but these authors agree that the fossils predate the emergence of the Isthmus of Panama. Overwater dispersal to South America by procyonids and sigmodontines is also covered by Koepfli et al. (2007) and Steppan et al. (2004), respectively. The radiation of South American sigmodontines described here refers only to the descendants of the presumed single oceanic dispersal event, that is, Oryzomyalia plus Neusticomys (Ichthyomyini). Neusticomys is placed in this group by Parada et al. (2013). Other lineages of sigmodontines, namely, the tribe Sigmodontini and some Ichthyomyini, may have reached South America separately and
perhaps by land over the Isthmus of Panama (Steppan et al. 2004).

  290Two much better known examples that fall into this category: The latest possible ages for colonizations by monkeys and caviomorphs are based on the earliest New World fossils for these groups (Takai et al. 2000; Antoine et al. 2012).

  291A great range of forms: The diversity of caviomorphs and sigmodontines is described in Lord (2007).

  291World’s largest known extinct rodent: Rinderknecht and Blanco (2008).

  293All three lineages reached the West Indies: For monkey dispersal to West Indies, see notes to Chapter Nine. For sigmodontine dispersals to West Indies, McFarlane et al. (2002); McFarlane and Lundberg (2002). For caviomorph dispersal to the West Indies, Pregill (1981). For sigmodontine dispersal to the Galápagos, Steadman and Ray (1982); Key and Heredia (1994). For sigmodontine dispersal to Fernando de Noronha, Carleton and Olson (1999).

  293Some that must indicate a massive impact: The numbers of species are from Lord (2007).

  293Tree of these nematodes largely mirrors: The nematode and monkey study is Hugot (1998). For some of the many parasites that are likely restricted to South American overwater colonist mammals, Tantaleán and Gozalo (1994); Rossin et al. (2010).

  294Snakes called the Xenodontinae: The time period for colonization of South America by xenodontine snakes is inferred from the timetree in Hedges et al. (2009; nodes 1 and 3 in Table 3), in conjunction with the larger phylogeny presented in Vidal et al. (2010a). The number of South American xenodontine species is from Vidal et al. (2010a), with the Alsophiini and other non–South American taxa removed. For xenodontine diversity and diet, Greene (1997).

  295The geographic origins of most South American plants: Renner (2004a) has compiled a list of at least 110 plant genera, including species in both tropical America and tropical Africa, most of which likely represent dispersal across the Atlantic either to or from the New World tropics. Molecular dating also has revealed many other cases of overwater dispersal by plants to South America (for instance, the tepui sundew and many of the bean-plant examples discussed in Chapter Seven). For recent reviews of the origins of Neotropical plants, see Pennington and Dick (2004); Christenhusz and Chase (2013).