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Australian Journal of Botany Volume 49, 2001 © CSIRO 2001

An international journal for the publication of original research in plant science All enquiries and manuscripts should be directed to: Australian Journal of Botany CSIRO Publishing PO Box 1139 (150 Oxford St) Collingwood, Vic. 3066, Australia Telephone: +61 3 9662 7613 Fax: +61 3 9662 7611 Email: [email protected] Published by CSIRO Publishing for CSIRO and the Australian Academy of Science

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Aust. J. Bot., 2001, 49, 389–409

Gondwana, vicariance biogeography and the New York School revisited G. NelsonA and P. Y. Ladiges School of Botany, The University of Melbourne, Vic. 3010, Australia. A Corresponding author; email: [email protected] Geography is impartial. Osmar White

Abstract. The many methods of biogeographic analysis proposed in recent years generate artefactual results that impede understanding, discovery and progress. Eliminating geographic paralogy from data reduces or eliminates artefactual interpretation. Recent cladistic studies of extant Nothofagus agree in showing only three informative nodes relevant to intercontinental relationships. In cladistic representations of global distributions, Gondwana is at or near the base of the geographically informative nodes, which force Gondwana to appear as a centre of origin of modern life in general. Centres of origin are artefacts of comparison based on geographically uninformative and paralogous nodes. Postmodern revivals of dispersalism fail to acknowledge, explain, avoid, learn from and improve on the artefactual centres of origin of the 20th century dispersalism, as represented particularly by the New York School: W. D. Matthew (1871–1930), K. P. Schmidt (1890–1957), G. G. Simpson (1902–1984), P. J. Darlington, Jr (1904–1983) and G. S. Myers (1905–1985). BGV.TicN0aeirsl02ao5nceabdnigoP.eYo.Lgardpigheysand theNew YorkScho l

Biogeography: a mess of methods Biogeography has been reviewed, more or less comprehensively, several times in recent years. The reviews testify to a continuing interest in their subject, stemming in the 1960s from the revival of continental-drift theory and from the development of cladistic systematics. From the decades since, the literature forms an interconnected whole, contemporary with the development, among other things, of postmodernism (Llorente Bousquets and Espinosa Organista 1991; Crisci and Morrone 1992; Espinosa Organista and Llorente Bousquets 1993; Morrone and Carpenter 1994; Reynoso Rosales 1994; Enghoff 1995; Morrone and Crisci 1995; Turner 1995; Morrone et al. 1996, 1997; Biondi 1998; Craw et al. 1999; Humphries and Parenti 1999; Vuilleumier 1999; Glaubrecht 1999–2000; Burckhardt and Basset 2000; Humphries 2000; Schuh 2000; Crisci in press; Crisci et al. 2000; Van Veller 2000; Ebach and Edgecombe in press; Espinosa et al. in press). At a recent international meeting (Anon. 1998: ix), there was a symposium ‘Historical biogeography: a critique’. The symposium provoked the response that ‘Biogeography is a mess’. As explained by its reporters, the meaning of the remark is that ‘biogeography is nowhere near finding a unifying and scientific method’ (Tassy and Deleporte 1999: 14, translated). The meaning is similar to that of observations from earlier times (Keast 1977: 249) that ‘some © CSIRO 2001

writers have recently been critical of the lack of a unified methodology; e.g. Vuilleumier 1975’; also Rosen (1978) on Vuilleumier (1978)—the centuries-old tension between ecological and historical approaches to biogeography. The varied possibilities to interpret data about taxa, their interrelationships and their geographic distributions, may be seen as different methods of analysis of data of these kinds (as in the summary diagram of Ebach and Edgecomb in press: fig. 9). Different methods, even applied to the same data, tend towards different results. In a historical sense, different and conflicting results—different histories— cannot all be true. At least some must be artefactual and, therefore, method-generated. At most, some only, or we are driven to the view of Samuel Clemens (1897: 699) that ‘the very ink with which all history is written is merely fluid prejudice’ (emphasis added). Unavoidable it might seem, when faced with different methods, is choice among them. So, too, are hopes that one method might be better than others and that choice might be guided by objective criteria. If one method were preferred by all concerned persons, then there would be consistent results derived from the same data. If, however, its results were artefactual to any degree, then a consistent method in general use would perpetuate the same problem, history as artefact if not prejudice, which would thereby have become obscured— swept under the rug of its apparent solution and the attendant forces of social conformity. Obscured, too, by the cloud of 10.1071/BT00025

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calumny likely raised around anyone using a method conflicting with any currently in fashion. So testifies the recent history of cladistic systematics (Deleporte and Lecointre 2000) and the emerging vanity that there be but one way, believed technically perfectible if not already perfect enough, to skin the cat of systematic endeavour (Platnick et al. 1997). On a broad scale, there is postmodern culture, itself a product of the 1960s according to one commentator, ‘characterised by continual shifting surfaces and a new ‘depthlessness’ derived from ‘the shock of slowly becoming aware that we are condemned to seek History superficially’ (Adams 1999: 107 after Jameson 1991). Maybe the problem, if there is one to address constructively, is only the need to find a way to better understanding. Geographic paralogy In 1996 we suggested that artefactual results stem from data that contain geographic paralogy (Nelson and Ladiges 1996). Use of the term ‘paralogy’ is by analogy with its use in molecular systematics begun by Walter Fitch in the late 1960s (Fitch 1970, 2000). There, paralogy refers to misleading comparison between duplicated genes that have had independent histories. Geographic paralogy is evidenced for different taxa by their partly or wholly overlapping distributions—duplicated geographies of taxa that have had independent genetic histories. These matters reduce to two novel ideas: (1) that a node of a cladogram, representing a phylogenetic relationship among organisms, is either geographically paralogous or it is not; (2) that any interpretation of a paralogous node is apt to be artefactual. We suggested that geographic data associated with non-paralogous nodes are the only such data actually relevant to cladistic biogeography. In our experience, removing paralogy from data prevents artefactual interpretation, if not altogether then to a significant degree. We expect that this will prove to be the case also in the experience of other persons (e.g. Anderson 1998) and time will tell. Nothofagus—southern beech trees For Darlington (1965: 140), Nothofagus was ‘the key to the history of terrestrial life in the far south’ because (p. 147) it ‘is likely to disclose a geographic history that has been followed by many other plants and by many invertebrate animals’ (also Van Steenis 1971, 1972, 1979; Veblen et al. 1996; Scriven 1997). Darlington (pp. 146, 147) explained: ‘Nothofagus may have originated in (southern?) Asia in the Cretaceous, crossed the tropics to Australia or New Zealand or both, radiated there and somehow made a triple dispersal half way around the southern end of the world’. This means three dispersals to explain the three subgroups recorded for South America (e.g. Van Steenis 1972: fig. 2; see also Takhtajan 1969: 154, 162; Whitmore 1981: 80; Hill 1992:

Fig. 1. (Left) Interrelationships of 30 extant species of Nothofagus, with their native area indicated: Aust, Australia; N Ca, New Caledonia; N Gu, New Guinea; NZ, New Zealand; S Am, South America, after Linder and Crisp (1995: fig. 3). (Right, above) The three geographically informative nodes (1–3) with relevant areas, derived from diagram at left. Nodes marked 0 are basal for paralogyfree subtrees (right). All other nodes are geographically paralogous. (Below) Combination of three informative nodes from above (Node 1 represents nodes 1+2 above, Nelson and Ladiges 1991: table 6, example 6). The tree (left) is stated as a strict consensus of 18 trees, but shows only one collapsed node, with a maximum of three possible resolutions. The tree is based on combined data: 26 morphological characters (Hill and Jordan 1993) and 46 informative characters from rbcL sequences (Fig. 2). Analysed separately by Linder and Crisp (1995: fig. 1), the morphological data yield a strict consensus of 12 trees; of 14 nodes, only one is geographically informative (equivalent to Node 1). For 31 species Hill and Jordan (1993: fig. 1; also Hill and Dettman 1996: fig. 2.1) found a strict consensus of four trees; of 19 nodes, three are geographically informative, equivalent to the combination (right, below) with S Am sister to NZ-Aust. Also on the basis of morphology, an earlier ‘tentative phylogeny’ shows two geographically informative nodes in one subtree: ((N Ca N Gu) NZ) S Am (Melville 1973: fig. 5, Cracraft 1975: fig. 5, Humphries 1981b: fig. 21.7). One of Humphries’ trees (fig. 21.9) has one geographically informative node: (N Ca N Gu) NZ Aust S Am and the other tree (fig. 21.10A) has none at all.

fig.1, 1994: fig. 16.1). The explanation implies that the South American representatives of each subgroup would prove related most closely to species of the jumping-off points for the three eastward dispersals. So far, this has proved not to be the case and is contradicted by subsequent discovery, as if the three hypothetical dispersals really had been from South America to the areas in the west (for ‘more than three’ see Fleming 1963: 379; Cracraft 1975; Melville 1982: 80). See for origin in North America and dispersal via Asia and/or South America (Oliver 1925; Schuster 1976);

Vicariance biogeography and the New York School

Fig. 2. (Left) Interrelationships of 23 extant species of Nothofagus, with their native area indicated (see Fig. 1), after Martin and Dowd (1993: fig. 2) and Manos (1997: fig. 2, in which N. aequilateralis is shown in place of N. codonandra and N. discoidea). (Right) The three geographically informative nodes (1–3) with relevant areas, derived from diagram at left. The tree is a strict consensus of 53 trees (according to Manos 1997: 1142), based on 46 informative characters from 1345 bp of chloroplast DNA rbcL. Linder and Crisp (1995: fig. 2) find a different strict consensus of 54 trees. Of 15 nodes, two are geographically informative (equivalent to Nodes 1 and 2).

Eurasia and dispersal via Africa–India (Raven and Axelrod 1972, 1974: 573); ‘an area of Gondwanaland which included Chile and Patagonia, Western Antarctica and the New Zealand platform’ (Melville 1973: 444); ‘a region between New Zealand, Antarctica and Australia’ (Hanks and Fairbrothers 1976: 69); Antarctica (Moore 1972; Dettman 1989, 1994; Dettman and Jarzen 1990; Dettman et al. 1990); ‘the region encompassed by southern South America and the Antarctic peninsula’ (Hill 1996: 247, Hill and Dettman 1996: 13; Hill et al. 1996: 182, 1999: 283; Veblen et al. 1996:

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Fig. 3. (Left) Interrelationships of 24 extant species of Nothofagus, with their native area indicated (see Fig. 1), after Setoguchi et al. (1997: fig. 3). (Right) The three geographically informative nodes (1–3) with relevant areas, derived from diagram at left. The tree is a strict consensus of 31 trees, based on 146 bp of rbcL and 453 bp of atpB-rbcL intergenic spacer.

388; Swenson et al. 2000a); a region ‘impossible to locate’ (Truswell et al. 1987: 44; Tyberg and Milberg 1998). In 1997 we considered the geography of Nothofagus, represented by 30 of the three dozen extant species in the strict consensus tree of Linder and Crisp (1995: fig. 3). Of 27 nodes in the tree, based on morphology and molecular sequences, only three are geographically informative (Fig. 1). Nodes 1 and 2 relate Australia and New Zealand more closely than to South America. Node 3 similarly relates New Caledonia and New Guinea (Ladiges et al. 1997: 128; Ladiges 1998: 236–237). Nothofagus is not richly informative about intercontinental relationships. The three nodes, nevertheless, are mutually consistent even if

Fig. 4. (Left) Interrelationships of 22 extant species of Nothofagus, with their native area indicated (see Fig. 1), after Manos (1997: fig. 5). (Right) The three geographically informative nodes (1–3) with relevant areas, derived from diagram at left. The tree is stated as a strict consensus of six trees, but shows only one collapsed node, with a maximum of three possible resolutions (one shown by Manos 1997: fig. 6; Swenson et al. 2000b: fig. 2, which includes two other New Caledonian species in place of N. aequilateralis). The tree is based on combined data: 1345 bp of rbcL (Fig. 2), 588 bp of ITS nrDNA and 23 morphological characters (modified from Hill and Jordan 1993). Analysed separately, the morphological data yield a strict consensus of eight trees (Manos 1997: fig. 3). Of 13 nodes, two are geographically informative (equivalent to Nodes 1 and 3). Manos (1997: fig. 7) included Nodes 1–3, each within its subtree, as one ‘area cladogram’ (7 nodes) in which areas appear more than once (Aust and NZ twice, SAm thrice). Subtrees 2 and 3 are shown as related (by Node 7) more closely than to subtree 1. Node 7, leading to the basal nodes of Subtrees 2 and 3, is paralogous and geographically uninformative.

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Fig. 5. (Left) Interrelationships of 22 extant species of Nothofagus, with their native area indicated (see Fig. 1), after Manos (1997: fig. 2). (Right) The three geographically informative nodes (1–3) with relevant areas, derived from diagram at left. The tree is a strict consensus of two trees, based on 588 bp of ITS nrDNA.

minimally corroborative: one node’s worth of corroboration. These results are similar to those, paralogy aside, of Humphries et al. (1986: fig. 4.14; cf. Patterson 1981 and Humphries 1981a, 1981b, 1983 for both of whom ‘Nothofagus is uninformative on the interrelationships of Southern Hemisphere areas’).

Fig. 7. (Left) Interrelationships of 22 extant, and from Tasmania (T) six fossil, species of Nothofagus, with their native area indicated (see Fig. 1), after Jordan and Hill (1999: fig. 4). (Right) The two geographically informative nodes (1–2) with relevant areas, derived from diagram at left. The tree is a strict consensus of 90 trees, based on combined data (Fig. 6).

G. Nelson and P. Y. Ladiges

Fig. 6. (Left) Interrelationships of 22 extant species of Nothofagus, with their native area indicated (see Fig. 1), after Jordan and Hill (1999: fig. 2). (Right) The three geographically informative nodes (1–3) with relevant areas, derived from diagram at left. The tree is a strict consensus of two trees, based on combined data: 24 morphological characters and 121 informative characters from sequences of rbcL (Fig. 2) and ITS (Fig. 5).

The molecular sequences are for 23 of the 30 species, for which Martin and Dowd (1993: fig. 2) published a strict consensus. Of 16 nodes in the tree, only three are geographically informative (Fig. 2). The three nodes have the same information as the three nodes of Linder and Crisp. For 24 species, including 22 of the 30, Setoguchi et al. (1997: fig. 3) published a strict consensus based on new molecular sequences. Of 15 nodes in the tree, only three are geographically informative (Fig. 3). The three nodes have the same information as the three nodes of Linder and Crisp. For 22 of the 30 species, Manos (1997: figs 2–6) published trees based on morphology and molecular sequences, some new. For the combined data, a strict consensus of 19 nodes (1997: fig. 5) has only three geographically informative nodes (Fig. 4). The three nodes have the same information as the three nodes of Linder and Crisp. For the new molecular sequences analysed separately, Manos (1997: fig. 2) published a strict consensus. Of 19 nodes in the tree, only three are geographically informative (Fig. 5). Node 2 differs from those above in relating Australia and South America more closely than to New Zealand. For a similar combined analysis of the 22 species, Jordan and Hill (1999: fig. 2) published a strict consensus (19 nodes), having three geographically informative nodes with the same information (Fig. 6). The addition of six fossil species from Tasmania yielded a strict consensus (1999: fig. 4, 17 nodes) having only two geographically informative nodes (Fig. 7), with the same information (Australia and New Zealand related more closely than to South America). The third node above, relating New Guinea and New

Vicariance biogeography and the New York School

Caledonia more closely than to South America, is rendered paralogous by the placement of two fossil species from Tasmania (N. mucronata, N. lobata), the former within a New Guinea–New Caledonia clade in an unresolved polytomy with six extant species, the latter within a resolved South America clade as sister to one of four extant species (N. nitida). Paralogy v. local precision One cause of paralogy is ‘imprecise characterisation of geographic areas’ (Nelson and Ladiges 1996: 11–12). Areas commonly used to describe intercontinental relationships of Nothofagus (South America, etc.) are not precise. Species of Nothofagus have more or less discrete distributions within each such area. More precise characterisation would doubtless expose certain nodes as non-paralogous, interrelating local areas within South America, New Zealand, Australia, New Guinea and even New Caledonia. Paralogy, corroboration, consistency and conflict By logical necessity, different subtrees imply (interconnect by) paralogous nodes. If informative of geographic relationship, different subtrees are corroborative, consistent or conflicting. Corroboration occurs when different subtrees include the same three (or more) areas interrelated in the same way by one (or more) non-paralogous node. Conflict occurs when such areas are interrelated in different ways. Consistency with no corroboration occurs when subtrees include fewer than three such areas and logically combine in a single and more informative tree. Without paralogy (in a general sense inclusive of molecular systematics), there is no possibility either of corroboration (Nelson 1994:138) or of conflict. Nothofagus, ratite birds and Gondwana Craw, Grehan and Heads recently contrasted Nothofagus and ratite birds – kiwis and their relatives (1999: 27; also Craw 1983, 1985; Heads 1985, Grehan 1988b; Page 1989): Distribution patterns of the ratite birds and the southern beeches are in no way biogeographically homologous or congruent, nor were they once members of a widespread, ancestral Gondwana biota as is often suggested in the biogeographical literature (e.g. Humphries 1981a; Patterson 1981). Nothofagus is a member of a non-Gondwanan trans-Pacific fagalean alliance; the ratites are a Gondwanic group centered on the South Atlantic and Indian Ocean basins. In the Southern Hemisphere these groups are geographically sympatric only in southwestern South America and in eastern Australasia where their tracks intersect. In their view, Gondwana is evidently understood to embrace taxa with distributions across the present Atlantic and Indian Oceans but not across the Pacific, as if all

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Gondwanan elements of South America today, like the rheas, relate to Africa rather than to Australasia (Lee et al. 1997). Gondwana—a node? Consider Croizat’s summary of global distribution (1958: 1018, fig. 259, reproduced in Nelson 1983: fig. 6, 1985: fig. 1; Craw 1984a: fig. 1, 1988: fig. 13.11, 1991: fig. 13; Craw and Weston 1984: fig. 1; Chiba 1987: 120; Grehan 1988a: fig. 3; Henderson 1989: fig. 15; Llorente Bousquets 1991; Espinosa Organista and Llorente Bousquets 1993: fig. 2.9; Chow et al. 1996: 267; Janvier 1996: fig. 1; Colacino 1997: fig. 1; Fortino and Morrone 1997: fig. 1; Crisp et al. 1999: fig. 75; Humphries and Parenti 1999: fig. 1.8; Morrone 2001: fig. 14; and redrawn in Brown and Gibson 1983: fig. 9.8; Crisci and Morrone 1989: fig. 4; Grehan 1991: fig. 8, 1993: Fig. 3, 1995: fig. 7; Brown and Lomolino 1998: fig. 12.3; Cox 1998: fig. 2; Craw et al. 1999: figs 6–12, 7–2; Morrone 2000: fig. 3). At the most general level, its cladistic representation shows five nodes: Atlantic, Indian, Pacific, boreal and austral with no relationships shown among the five (Nelson 1985: fig. 1; Chow et al. 1996: 267). The last two, boreal and austral, have long been associated as bipolarity (Berg 1933; Du Rietz 1940; Andriashev 1987), which would form an additional, sixth, node to the cladistic representation (Fig. 8A, B). There is reason to associate bipolarity with the Pacific rather than with the Atlantic and Indian Oceans and this association would form an additional, seventh, node (Fig. 8A, B), the ‘Pacific Rim’ of Craw et al. (1999: 27). Bipolarity, even extended to the Pacific Rim and beyond to the Caribbean (Rosen 1976; cf. Briggs 1994), does not exhaust the complexity of Pacific distribution (Van Der Spoel 1983; Nelson 1986; Eskov and Golovatch 1986; Sluys 1994; Shields 1998; Heads 1999). To the cladistic representation in our view, Craw et al. would add an eighth, and last, node, associating the Atlantic and Indian Oceans (Fig. 8A). To that node they would restrict

Fig. 8. (A, B) Cladistic representations of Croizat’s (1958: fig. 259) summary of global patterns of distribution of plants and animals. Nodes: 0, uninformative; 6, bipolar; 7, Pacific Rim; 8, Gondwana according to Craw et al. (1999), with alternative placement shown in B.

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the meaning of the term ‘Gondwana’. With respect to ratites and Nothofagus, their evidence is non-existent. Ratites might be Atlantic and Indian and Nothofagus Pacific. It does not follow that the two (Atlantic, Indian) relate more closely than to the one (Pacific). Craw et al. note (p. 26) that ‘Nothofagus is unknown as an autochthonous fossil from Africa and India (Hill 1994), the heart of the Gondwana supercontinent.’ Nothofagus forests might be unknown from the heart of Gondwana, but they amply drape over her left arm and shoulder. Bony-tongue fishes and Gondwana We refer here to South America and two extant freshwater fishes of the bony-tongue family, Osteoglossidae: the arawana (Osteoglossum) and the pirarucú (Arapaima) of Amazonian white water (Fig. 9). They live together over a vast distribution (Berra 1981). It is said that the smaller arawana is the ‘favourite bait’ of the larger pirarucú (Herald 1961) or is sometimes eaten (Roberts 1972), but is not always recorded as an item of the natural diet (Lüling 1964, 1971). The pirarucú, one of the world’s giant fishes, has its nearest living relative in Africa (Heterotis, tropical West Africa and the upper Nile; Roberts 1975). The arawana has its nearest relatives in a complex of species (Scleropages): one or more species in Sumatra, Borneo and South East Asia from Malaya into Thailand; two or more species in New Guinea and tropical Australia, which have the name saratoga in recent Australian books (Merrick and Schmida 1984; Grant 1987; Larson and Martin 1990) and burramundi or barramundi in earlier books (Whitley 1960; Munro 1967; Lake 1971, 1978; Pollard 1980), a name now reserved in commerce for an unrelated fish (Anon. 1995; Yearsley et al. 1999). The New Guinea and Australian forms are related more closely than to the Malayan species (on the basis of personal observations). Possibly they are more closely related to the South American arawana than to the Malayan species (Nelson 1969a). More remote are relatives, fossil in India (Bonde 1996) and extant and fossil in Africa (Li and Wilson 1996: fig. 4). The relevance of the fishes is the evident trans-Pacific relationship within a group, evidently Gondwanan, with fossil roots into the mid-Mesozoic (Kumazawa and Yoshida 2000). In terms of their geography, the fishes are nearly equivalent to the ratites and Nothofagus combined, with a trans-Pacific distribution extending beyond the shoulder into India and Africa, the heart of Gondwana. So in South America is the pirarucú a Gondwanan element and the arawana not, because the one relates to the east and the other to the west? Trans-Pacific relationships, Gondwana and the world Craw et al. (1999: 154–160) consider Pacific distribution and Croizat’s (1960: fig. 8) concept of a geologically composite New World (reproduced in Craw 1984a: fig. 2,

Fig. 9. Interrelationships and distribution of extant fishes of the family Osteoglossidae, including three species of Scleropages, with their native area indicated (see Fig. 1 and text). Nodes: 0, Paralogous; 1, Atlantic; 2, Gondwana; 3, Pacific; 4, Arafura Sea— Torres Strait.

1984b: fig. 4, 1991: fig. 7; Craw and Weston 1984: fig. 3; Grehan 1988a: fig. 6; and redrawn in Craw and Page 1988: fig. 9; Grehan 1991: fig. 18, 1994: fig. 3; Cox 1998: fig. 1; Craw et al. 1999: figs 6, 7; Morrone 2000: fig. 4). In Croizat’s view, the western sector, from Alaska to Chile, relates to lands bordering the West Pacific, but not in their view to Gondwana. Yet relevant lands of the West Pacific are seen as physical parts of a geologic Gondwana (Metcalfe 1999; Metcalfe et al. 1999; Michaux and White 1999). So, too, are Pacific terranes—harbouring trans-Pacific biological relationships—seen as physical parts of a geologic Pacifica continent and of a more inclusive Proto-Gondwana landmass (Craw 1985: fig. 5, 1988: fig. 13.9, 1991: fig. 5; Chow et al. 1996: 253). Are trans-Pacific biological relationships external to or embedded within Gondwana and her history? As a biogeographic relationship of today, is Gondwana to be understood to relate distributions across the Atlantic and those across the Indian Oceans? Or, as an alternative and informative Node 8, to relate distributions over one or the other possible combination of oceans, Indo-Pacific or Atlanto-Pacific? In Croizat’s summary, the only other node available for the name ‘Gondwana’ is the cladistically uninformative basal node of life’s global distribution (Fig. 8B). In that sense Gondwana and today’s world would be one. This alternative implies that the life of the past Gondwana is reflected today in the distribution of life the world over, a fair summary of the actuality. For a modern distribution to appear Gondwanan, rather than merely Atlantic, Indian or Pacific, there seem to be two requirements: that it involve more than one ocean basin; and that in a phylogenetic tree it associate with a node, likely to be geographically paralogous, that is basal enough so that all more basal nodes are also paralogous. Such nodes also invite association with Gondwana. These requirements (and nodes) force Gondwana to appear, artefactually, as an ultimate centre of origin of modern life in general (e.g. Cracraft 2001).

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Centre of origin? Faced with paralogy, various notions self-destruct. One notion, perhaps the most notable casualty, is again, the centre of origin, which in a cladistic context seems always a product of paralogous comparison (Ebach 1999). Consider a hypothetical example (Fig. 10), sent to Willi Hennig a few years before he died in 1976. There are two taxa (extant or fossil) in South America, one related to a third in Africa (Nelson 1974: fig. 2). The question posed to Hennig was: is there evidence, however slight, of a centre of origin in South America and dispersal to Africa? Yes, according to his Progression Rule (Hennig 1950: 356, 1960: 250, 1966a: 134, 1966b: 23, 1968: 183, 1982: 132; Brundin 1966: 56)—the usual cladistic criterion (Fig. 11; Ross 1974: 214–220; Platnick 1981; Wiley 1981; Nelson and Platnick 1984a); no, if the basal node be set aside as paralogous. Is it reasonable in this hypothetical case, despite the duplicated geography in South America, to consider that the ancestor of the three taxa occurred also in Africa? Yes, why not? The cladistic criterion used to resolve a centre of origin is minimality of implied dispersal events (Nelson 1969b). Why not minimise them to zero? Without a centre of origin and dispersal therefrom, what is the implied history in this hypothetical case? That the first split of the widespread ancestor isolated a population in South America before the later split between the less widespread South American–African population.

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interpret geography to fit paralogous nodes in the hope that a meaningful pattern might emerge from their analysis even by a computer program. Geographic homoplasy? A third notion is that apparently conflicting and artefactual geographic data are analogous to homoplasy in systematic data, which, hopefully, is distributed randomly, more or less, among the nodes of a cladogram—not so because geographically paralogous nodes increase towards the base of cladograms generally. Consider two nodes: one node (non-paralogous) relating a pair of closely related fishes, or a pair or aquatic insects, living in adjacent river basins, and another node (paralogous) deeper in the history of life, relating fishes and the insects upon which they feed. The former is geographically relevant and the latter not. The former node speaks to a relationship between the river basins. The latter to who knows what? Creation myth: past and present

A second notion is that attached to each node of a cladogram there are relevant geographic data deserving of interpretation —not so for paralogous nodes. There is no necessity to

A remarkably recurrent theme of 20th century biogeography is the quest for origins of taxa and of their distributions, or in other words ‘The geographic origins of individual taxa and dispersal routes away from these origins’ (Simberloff 1972: 161), the ‘point of origin (evolution/migration)’ of Macphail (2000a: 19). The notion of a geographic centre of origin, with dispersal therefrom, explicitly derives from the creation myth of the Bible – the Garden of Eden. Linnaeus and later writers used the myth as an explanatory device of biogeography (Nelson and Platnick 1981; Nelson 1983). It persists in the ‘out of Africa’ sagas of human origins and their molecular embellishments. It is currently under postmodern revival: if the flora of New Zealand, ‘isolated

Fig. 10. In South America, two taxa (extant or fossil) of which one relates to a third in Africa, showing possibilities for vicariance (left) and dispersal (right).

Fig. 11. (Top) In South America, two taxa (extant or fossil) of which one relates to a third in Africa, illustrating minimum evidence (paralogous Node 1) to resolve a centre of origin according to the usual cladistic criterion (Hennig’s Progression Rule). (Middle) Twice the minimum evidence (paralogous nodes 1–2). (Bottom) Thrice the minimum evidence (paralogous nodes 1–3). Simpson (1947: 649) regarded such evidence (‘a variety of earlier relations’) as ‘more reliable than…earlier appearance’ in the fossil record.

Paralogous nodes—geographically informative?

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from Australia and Antarctica...for 60 million years...can be shown to have arrived over the sea...then biogeographic hypotheses the world over which involve any kind of land connection must be reconsidered’ (Pole 1994: 625). The argument is the familiar ‘if-then’ of the ‘professional biogeographer’ (Darlington 1964a: 1084): ‘if Nothofagus has been wind-dispersed across southern ocean gaps in the late Cretaceous and Tertiary, [then] Glossopteris may have been dispersed in the same way in the late Paleozoic’ (Darlington 1965: 147–148). With an explanatory mix of dispersal and vicariance, determined by ‘vastly improved geological data’, biogeography is now seen suddenly to have ‘got real’ (Heaney 1999: 435–436): For me, for example, knowing with good confidence that the continental rocks included in the Philippine island of Mindoro were covered by marine seas at the time of their fragmentation from the Asian continent and that they did not re-emerge as part of a subaerial island until the late Miocene (roughly 10 million years ago) makes all the difference in successfully sorting process out of patterns of distributions. The possibility of complete submergence of New Zealand in the late Oligocene and of New Caledonia in the early Tertiary, is similarly used to argue for ‘entirely long-distance dispersal’ as explanation of the biota of these islands (Pole 1994: 628–629). Once again (Darwin 1845: 378) ‘we seem to be brought somewhat near that great fact—that mystery of mysteries’…the vision that ‘the floras and faunas of many islands, including New Zealand...must have crossed water gaps’ (Darlington 1964c: 708; Macphail et al. 1994; Macphail 1997a, 2000b; Swenson and Bremer 1997; Wagstaff and Garnock-Jones 1998; Winkworth et al. 1999). Familiar, too, are the suggested means of this long-distance dispersal (McGlone et al. 1996: 83)—the ‘flotsam of sea and sky’ (Cranwell 1963: 387): ...obvious candidates are storm-force winds and the feet and plumage of birds. While the probability of this happening in any one year is likely to be extremely small, migrational lags of millions of years may have provided the requisite time interval for the improbable to happen. (cf. Matthew 1915: 206–209, 1939: 37–40; Simpson 1952; Darlington 1957: 484). Emerging lifeless from the ocean, barren lands feature in Linnaeus’ explanation of life’s distribution throughout the world (Linnaeus 1744, 1781; Browne 1983; Frängsmyr 1983; Seberg 1985; Larson 1986, 1994; Rupke 1996, 1997; Papavero et al. 1997; Bueno et al. 1999; Llorente Bousquets et al. 2000; Bueno Hernández and Llorente Bousquets 2001). Whether universally or locally applied, the vision and its effect are the same—to render superfluous any geographic comparison across taxa. Biogeography thereby reduces to dispersalism and its empirical impossibilities—marking

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the centre of origin and time of arrival of each, or even one, propagule theorised to have been successful in colonising the barren landscape to the east, or to the west, of Eden. To all appearances, a real problem (what to believe? or how to proceed? or when and where to stop?) is thereby solved. That the history of biogeography is littered with solutions of the kind testifies to their continuing popularity (Nelson 1978). Popular, too (Macphail 1997b: 425), is the modern resonance with the classical dictum, from Aristotle (Historia Animalium, book viii, s28) via Pliny (Naturalis Historia, book viii, s17), that ‘out of Africa there is always something new’—Ex Africa semper aliquid novi (Smith 1939). The modern resonance stems from Darwin’s (1871: 199) comments on the ‘Birthplace and Antiquity of Man’: It is therefore probable that Africa was formerly inhabited by extinct apes closely allied to the gorilla and chimpanzee; and as these two species are now man’s nearest allies, it is somewhat more probable that our early progenitors lived on the African continent than elsewhere. The logical principle of Darwin’s ‘cogent reasoning’ (Leakey 1960: 18) later became generalised as the progression rule to circumscribe a centre of origin from phylogenetic trees, including those based on fossil as well as molecular data (Stoneking 1996: fig. 11.4; Patterson 1999: fig. 16.1). The principle underpins the answer to the question, ‘Where are we to pitch the centre of dispersal? The evidence, as it stands today, favours Africa…If we may select one region as more likely than another, then our choice falls on the uplands of Uganda and Kenya’ (Keith 1948: 214). The evidence is paralogous nodes, or the equivalent nodes of presumably direct ancestry (Nelson 1973: fig. 1). Recast by Louis Leakey as ‘Charles Darwin’s prophecy’ (also Tobias 1984: 37), the modern resonance peaked in a series of fossil finds (Tattersall 1995), beginning most notably in 1924 with (Leakey 1974: 193) ‘Raymond Dart’s spectacular discovery of the juvenile skull of Australopithecus at Taung, South Africa’: I stood in the shade holding the [fossil] as greedily as any miser hugs his gold, my mind racing ahead. Here, I was certain, was one of the most significant finds ever made in the history of anthropology. Darwin’s largely discredited theory that man’s early progenitors probably lived in Africa came back to me. Was I to be the instrument by which his ‘missing link’ was found? (Dart 1959: 6). Soon, it was said that ‘the combined…efforts of Dart, Broom, Leakey and Oakley have established the rough but indisputable outline of the human emergence on the African highland’ (Ardrey 1961: 27). So developed the story of ‘that great fact—that mystery of mysteries—the first appearance of new beings on this earth’ (Darwin 1845: 378). Associated with Louis Leakey (1903–1972) is another prophecy, which he did not survive to judge at the stated time

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—that the future, even 10 years hence, would see ‘how little we knew, how stupid we were in 1965!’. His doubt extended even to the ideas that the human family ‘Hominoidea started in Africa’ and, accordingly, that ‘there were…movements out into Europe and Asia’ (Leakey 1965: 17, 1972: 399). By 1969, however, Leakey’s doubt was seemingly forgotten (Leakey and Goodall 1969: 170): Charles Darwin’s prophecy is coming true. More and more evidence is accumulating which points to the African continent and particularly the East/Central African region, as the cradle of the Family Hominidae, to which all mankind, living and extinct, belongs. And so began the recitations of the ‘journey from Eden’ (Fagan 1990), the ‘African exodus’ (Stringer and McKie 1996), the ‘footsteps of Eve’ (Berger 2000), the ‘Genesis chronicles’ (McBride 2000). The question naturally arises (Foucault 1977: 144): ‘does this not form a history, the history of an error that we call truth?’. New York School Of dispersalism, its history in the century past belongs also to what Croizat (1958: xi) called the ‘New York School of Zoogeography’, in reaction to which vicariance biogeography developed (Nelson and Rosen 1981). Among New Yorkers, the principal players in the ‘great and splendid drama’ are fossil vertebrates, mammals in this case and persons posing as their interpreters: primarily William Diller Matthew (1928: 54; Bowler 1996) and secondarily George Gaylord Simpson. In their view the crucial evidence is provided by discovery of the fossil ancestors of a taxon— plesiomorphic fossils. Once found, fossil ancestors—the ‘true genetic sequences’ (Matthew 1925: 288)—reveal directly the centre or place of origin of the taxon in question: the geographic space inhabited by the ancestors. That this empirical discovery was possible, even ‘demonstrably true’ (Simpson 1937: 253), they took for granted and never questioned. When occasionally this empirical discovery seemed not forthcoming, they ‘somewhat sadly’ lamented its absence (Simpson 1978b: 324). It was early questioned during the development of cladistics and found impossible to achieve. Years of argument and discussion of the matter were later summarised by Colin Patterson (Fortey 1999; Forey et al. 2000) with his usual clarity and eloquence: ‘Fossils may tell us many things but one thing they can never disclose is whether they were ancestors of anything else’ (Patterson 1978: 133, 1999: 109). W. D. Matthew (1871–1930) Of Matthew it is said that ‘the stray lock of hair and the almost inevitable cigar were trademarks of the man’ (Colbert 1992: fig. 40). So wrote Ned Colbert, who in 1933 married William Diller’s younger daughter Margaret. Matthew wrote a celebrated essay of 150 pages, ‘Climate and Evolution’. Thirty years later, Karl Schmidt (1943: 242) wrote that the

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essay had inaugurated ‘a new phase in the study of animal geography’. Thirty years later still, Simpson (1978a: 272) described it as ‘one of the most seminal or heuristic studies of paleogeography and historical biogeography’. The essay was published in 1915 by the New York Academy of Sciences (Matthew 1915). Nearly 25 years later it was reprinted in 1939 (Matthew 1939) because, according to Colbert (1939: v) at the time, ‘there has been a steady demand for this paper that has continued to the present day, long after the original edition had been exhausted’. The essay was reprinted again in 1950 and, incredible as it may seem, reprinted again in 1974. For life generally, Matthew advocated northern (Holarctic) centres of origin with independent dispersal southward: to Africa, through South East Asia to Australia and across a Bering land bridge to and through North America to South America. His is the same notion that originated in the mid-19th century if not earlier. Ernst Haeckel (1834–1919) used it to explain the history of humans and their wanderings from their centre of origin in Paradise (Haeckel 1889: pl. 20 and later editions, modified from Haeckel 1879: pl. 15 and earlier editions beginning with the second [the many English translations begin with Haeckel 1876, French with Haeckel 1874], Nelson 1983: figs 1, 2; Patterson 1999: fig. 16.2; Thomas 1994: 40–41; Kirchengast 1998: fig. 2). With this equation of notions, Matthew might have disagreed (also Savage 1958: 154), pointing out correctly that his centres of origin are north of the languid tropics. In his centres, ‘greater activity and higher development of life’ would emerge in reaction to ‘the inclemency of nature, the scarcity of food, the variations of temperature, as well as against the competition of rivals and the attacks of enemies’ (1915: 177, 1939: 7). But Matthew, too, considered his (Matthew 1928: 81) ‘an old view which had been outlined first by Buffon [1707–1788] and was elaborated by [Alfred Russel] Wallace [1823–1913] in his great book on [The Geographical] Distribution of Animals [1876]’ (noted also in Croizat 1958: i). K. P. Schmidt (1890–1957) Matthew was seen as an inspiring adversary of continental drift. In the words of one of his students: In the very year when [Alfred] Wegener proposed his theory [Wegener 1915], this...was shown by William Diller Matthew to be invalid. So wrote Karl Patterson Schmidt (1955: 777), herpetologist at the Field Museum of Natural History, Chicago. He died in 1957, after being bitten on the tip of the thumb by a small snake brought to the Museum for identification from the local zoo (Anon. 1957; Davis 1959). He wrote also (pp. 780–781): A group of students in zoology and paleontology under William King Gregory at the American Museum of

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Natural History in the years 1915 to 1927 came also under the influence of W. D. Matthew...The group included Alfred Sherwood Romer, Charles Lewis Camp and Gladwyn Kingsley Noble, with many others and a little more indirectly Emmett Reid Dunn and myself. All of us have remained disciples of Matthew...we have tended to look a bit askance at those ‘who knew not Matthew’; and it had not escaped some of our colleagues that Matthew’s work had become a kind of Holy Writ to his disciples...The leadership among the group of Matthewsians has now somewhat naturally fallen to George Gaylord Simpson, who succeeded Matthew in the position of Curator in charge of Vertebrate Paleontology at the American Museum in 1944. G. G. Simpson (1902–1984) In 1924 while a graduate student at Yale University, the young Simpson was hired as assistant to Matthew, spending 6 weeks with him in field work in Texas (Laporte 1986, 1990). In 1927 Matthew left New York City for the University of California at Berkeley, there to found ‘the only separate department of paleontology at an American university’ (Colbert 1989: 143). Simpson replaced him in employment at the American Museum of Natural History (on Matthew’s recommendation) and in 1945 at Columbia University (Hecht et al. 1972; Laporte 1991). Simpson remarked: ‘I came to the American Museum to follow, at a great distance, in the footsteps of W. D. Matthew’ (Simpson 1945: vii). ‘He was a great paleomammalogist, a hero to me and with him familiarity bred only respect and admiration’ (Simpson 1984: xvi). ‘I absorbed every word he spoke and read...every word he had written’ (Rainger 1991: 213). ‘I soon was drawn to the essentials of W. D. Matthew’s views’ (Simpson 1976: 8). And with Simpson, as early as 1940, it was more of the same: ‘the general type of geographic history assumed by Matthew to be typical for mammals is...here more explicitly supported’ (Simpson 1940a: 141). ‘Matthew’s theory does afford the best explanation so far proposed’ (Simpson 1940b: 765). ‘The distribution of mammals definitely supports the hypothesis that continents were essentially stable throughout the whole time involved in mammalian history’ (Simpson 1943: 29). Simpson, too, eventually wrote an essay, of a modest and less celebrated 64 pages, ‘Evolution and Geography’, first published in 1953, reprinted a half-dozen times by 1968 and also translated into Spanish (Simpson 1964) and French (Simpson 1969). It concludes: ‘All the biogeographic features in the known history of mammals are best accounted for on the theory that the continents have had their present identities and positions...the conclusion seems to apply not only to the biogeography of mammals but also to that of all contemporaneous forms of life’ (Simpson 1953: 61–63, 1964: 55, 56, 1965: 130, 1969: 106, 107).

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After moving from the American Museum to the Museum of Comparative Zoology (MCZ) at Harvard University, Simpson remarked (1961: 1684): ‘Matthew...has successors who have followed in his footsteps and have, with constantly improved data, gone well beyond him. The main outlines...of mammalian faunal evolution are now well established.... Currently accepted general principles of historical biogeography...are derived largely from paleomammalogy in the tradition of Darwin and Matthew. This is evident, for instance, in a fine recent treatise...written by an [MCZ] entomologist, [Philip Jackson] Darlington [1957].’ Twenty years later Simpson (1980: 253) described this as an ‘excellent...treatise...on biogeography that is now out of date but better than most of the recent works on that subject’—the review by Keast (1977) being ‘incomparably the best modern summary of the whole subject’ (Simpson 1978c: 219). P. J. Darlington Jr (1904–1983) Other forms of life is the theme previously taken up by Darlington in papers featured in the Quarterly Review of Biology in 1938 and 1948. The former considers dispersal of organisms to the Caribbean islands by means of wind, cyclonic storms (hurricanes), waterspouts and ocean currents, and finds (p. 197): The fauna is…very orderly…[The] animals are still distributed along the migration routes by which their ancestors reached the islands and this is taken to show that the fauna is an accumulation of immigrants derived from the mainland across water…No other hypothesis will fit the facts. For George Myers (1954a: 19) the former was ‘an extremely important paper on the origin of the Greater Antillean fauna, in which [Darlington] concluded that [that] fauna gives no indication of a continental connection’. The latter paper, ‘The Geographical Distribution of ColdBlooded Vertebrates’, prompted Karl Schmidt to remark (1955: 780): Fortunately we have now had a strong cross-light thrown on the main thesis of Climate and Evolution by a nondisciple [of Matthew], P. J. Darlington, Jr, of the Museum of Comparative Zoology at Harvard University, who reviews the whole matter from the evidence of the freshwater fishes, amphibians and reptiles. Darlington’s approach was later perfected in another celebrated work. Considered ‘the most meritorious work in zoology published during the year’, the book ‘Zoogeography’ (Darlington 1957) was awarded the 39th Daniel Giraud Elliot Medal of the National Academy of Sciences of the United States (NAS Website 2000; Carpenter 1985: 10, but in the year 1957 not 1969). Not merely for cold-blooded vertebrates but also for birds and mammals, this book summarised the facts as seen from the viewpoint, thereby reaching its climax at Harvard University, of the

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New York School near the end of its development. The book marked the ‘end of an era’ (Savage 1958: 155), but not the end of that school’s influence, which, ‘in a virtually unassailable position’ (Simpson 1958: 379), continues to the present, ‘hardened into solid theory’ (Brown 1985: 15), most notably among ichthyologists (Banarescu 1970, 1990–1995, 1996; Banarescu and Boscaiu 1973, 1978; Briggs 1974, 1984, 1987a, 1987b, 1992, 1995a, 1995b, 1996, 1999a, 1999b, 1999c, 2000). In sum for Darlington (1959a: 313— ‘an article written to be read, not just filed in historical archives’ [Darlington 1980: 11], 1964b: 970): Darwin considered the evidence he had and decided that as far back as he could see the main pattern of land had been the same as now...Fifty-six years later Matthew, with much more evidence, reached the same conclusion, but saw farther back and in much more detail than Darwin could. And now, with still more evidence, we can see still farther back and in still more detail than Matthew could, but the conclusion is still the same. The existing pattern has evidently been formed by very complex movements of animals over the world approximately as it is now, not over extraordinary land bridges or drifting continents. G. S. Myers (1905–1985) Central to the ichthyological history, an academic conduit from New York to Cambridge and to the world at large— even to New Zealand (McDowall 1988: vii)—is George Sprague Myers and his ‘more than 104 graduate and special students’ at Stanford University, during his 35-year tenure as Professor in the Department of Biological Sciences (1936– 1970, Herald 1970). Earlier (1922–1924), Myers had been a volunteer assistant at the American Museum of Natural History. It is said (Walford 1970: 3, Cox 1988: 3) that he there came under the influence, among other persons, of Karl Schmidt, himself employed intermittently at that institution, eventually resigning as Assistant Curator in 1922 ‘to accept a position in Chicago as the first head of a new Division of Amphibians and Reptiles at the Field Museum of Natural History’ (Myers 2000). Before Matthew’s death in 1930, Myers (1938: 341) had apparently ‘gathered the ichthyological information’ relevant to ‘the proposed enlarged and revised [second] edition of ‘Climate and Evolution’.’ When that second edition was published (Matthew 1939), it was without his contribution, which had appeared shortly before (Myers 1938). Therein he divided true freshwater fishes into primary and secondary classes. More numerous in species, the primary freshwater fishes ‘possess an ancient physiological inability to survive in salt sea water, which binds them to the land as securely as any known animals’. Most primary fishes are in the group Ostariophysi, which include carps, characins, catfishes and electric eels.

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Secondary freshwater fishes ‘are generally restricted to fresh water but occasionally enter the sea’ (Myers 1938: 342–344). Other ‘still more salt-tolerant’ fishes found in freshwater he later grouped into four additonal classes (Myers 1949, 1954a, 1954b; Kessel 1970 explains dates of Myers’ publications). He stated that his accounts of the primary fishes (1938: 354): ...make it clear that our present knowledge of the fishes very distinctly favors a late Mesozoic or very early Tertiary South Atlantic land connection and makes a direct northern origin...seem exceedingly unlikely. But I refuse to take a definite stand on these questions. It would be extremely presumptuous, on the basis of the fishes alone, to attempt a flat contradiction of the Holarctic dispersal of the mammals, reptiles and amphibians so ably advocated by Matthew (1915), Noble (1925) and Dunn (1923 and 1931). In 1938 Myers (p. 341) wrote that ‘Climate and Evolution’ has had a profound and overwhelming effect on nearly all recent American zoogeographers and but little on anybody else’. By 1946, ‘after eight years’ cogitation on fresh-water fish dispersal’ Myers had ‘gone over to the northern origin idea, with all the speed and grace of a cat dragged backward by the tail over the dining room rug. There is simply no other answer to the fish distribution problem and I say this with perhaps fuller knowledge of the objections than anyone’ (Myers in litt. to Darlington, 12 November 1946). In 1949, he considered Matthew’s 1915 essay as ‘the most important single contribution to zoogeography since the time of Wallace’ (Myers 1954a: 19), a judgment surpassed only by Darlington (1959b: 488): after ‘the two chapters on geographical distribution and…parts of other chapters’ in the ‘Origin of Species’ (1859) ‘the next really important treatment of the subject was by Matthew (1915) in ‘Climate and Evolution’.’ Of Myers’ divisions into primary and secondary classes, he (1949: 317) noted that ‘Darlington has utilised these divisions in an extensively well-thought-out general scheme of fresh-water fish distribution’. He noted that ‘in 1948, Dr Darlington gave an excellent resumé of the world’s freshwater fish distribution, in which he expanded the fish data given in my 1938 paper and included amphibians and reptiles’ (Myers 1951: 11, 1954b: 39). For his part Darlington (1948: 4), as he had done 10 years before (1938: 291), acknowledged that ‘I am greatly indebted to Dr George S. Myers for guidance in my work on fishes...’. Of his divisions Myers wrote to Darlington that ‘I have made one discovery worth general quoting in my life, which I am human enough to want to see recognised. You did.’ (in litt. to Darlington, 17 April 1951). Of Myers’ 1938 paper it was noted (Walford 1970: 12) that ‘20 years after publication this paper was acknowledged by P. J. Darlington in his great book ‘Zoogeography’ as the prime reference on which he built that part of his book

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dealing with fishes’. For his part Darlington (1957: x–xi) conceded that ‘Thanks to hard work on my part and especially to aid from George S. Myers and other ichthyologists, what I shall have to say about fresh-water fishes is...perhaps better than any other part of this book’. By the early 1960s, Myers (1963; Kessel: 1970: 51) took the definite stand long deferred (anticipated in 1938: 340, 341), ‘deserting the Gospel according to St. Matthew’ (in litt. to Darlington, 21 January 1964) – rejecting notions of continental stability in favour of drift as explanation of geographic distribution of fishes: ‘I believe continental drift to be the ultimate answer and even Darlington [1964a] has very recently cautiously come out in favour of drift’ (Myers 1966: 772; for Darlington 1965 see Myers 1967: 620 and Hershkovitz 1972: 316). Cautiously indeed. According to Doctor Darlington the patient might be pregnant, but only a little bit (1964a: 1090, 1979: 345): Africa and South America probably were united but...separated not later than the Triassic and perhaps earlier, so long ago that no clear traces of the union are visible in distribution of existing life. Other southern continents were probably not united. Myers rejected continental stability because, with increased knowledge of Central American fishes (or increased meditation upon their significance), he again found implausible the Matthew–Darlington explanation of the primary freshwater fishes of South America – their dispersal through North and Central America from an Asian centre of origin. Ironically, this rejected explanation, so clear and comprehensive, was due partly to Myers’ own prior effort and history of shifting conviction (Walford 1970: 12). It is said (Géry 1969: 833) that ‘Darlington’s (1957) hypothesis...expressed rather faithfully the views of specialists with G. S. Myers at the head...’. And only 10 years before according to Schmidt (1955: 780): The very important evidence of the freshwater fishes, long regarded as a proof of the necessity for past direct connection of Africa and South America, is now regarded by George S. Myers as explainable by longterm round-about emigration via the Bering Bridge [and Central America] rather than by hypothetical trans-Atlantic connections. In the 1960s Myers did not adopt a new and alternative explanation, but rather the old one that by the mid 1940s he had previously ‘discarded’ (Myers 1951: 11, 1954b: 38). For fishes, the old explanation had been best developed by Charles Tate Regan (1878–1943) at the British Museum (Natural History) in the year (1922) that Myers began his volunteer work at the American Museum of Natural History, his physical entry into the New York School. At this time he presumably began to learn of ‘the strength mustered against them’ (Myers 1951: 11, 1954b: 38). This ‘strength’ he portrayed as a list of references: Matthew 1915; Noble 1925; Dunn 1931; Schmidt 1943; Simpson 1939, 1940a, 1940b;

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Darlington 1948; Berry 1928; Chaney 1940. ‘Them’ means persons—such as Regan, destined in 1927 to become the sixth Director of the museum in South Kensington (Burne and Norman 1943)—who favoured, among other explanations for various organisms, the old explanation for fishes: ‘that in early Cretaceous times S. America and Africa formed one continent, which must have extended to India...The alternative view, that the Ostariophysi originated in the north and spread southwards, involves so many improbabilities as to be almost unbelievable.’ (Regan 1922: 206–207). Using the phrase ‘strictly fresh-water’, Regan had considered the same fishes (Ostariophysi) under the same point of view later adopted by Myers, who used the term ‘primary’ rather than ‘strictly’. Regan’s (1922) paper was later promoted as an ‘antidote to this influence’ [of the New York School] by one of Myers’ own students (Gosline 1944: 213) in a paper that Myers (1949: 317) once cited as an example of what might appear to the reader as a minor issue: ‘doubt...as to the real distinctiveness of the two divisions’ (Myers’ primary and secondary divisions)—doubt as to the assumed difference in the fishes’ physiological tolerance to dissolved salt (also Gosline 1975). A difference in physiological tolerance is fundamental to what Myers considered his ‘one discovery worth general quoting’ (see above). The discovery stemmed, seemingly, from experience in keeping aquarium fishes. It existed also as hope and expectation that were never realised: ‘I hope (and rather expect) that…physiological work…will demonstrate a real basis for the primary and secondary divisions’ (in litt. to Darlington, 14 July 1948). For him a real basis would explain ‘why [primary fishes] are confined to fresh water’ (Myers 1949: 318) and justify the claim, or discovery, of their ‘ancient physiological inability to survive in salt sea water’ (see above). In his view this real basis, once demonstrated, would lie within physiological ecology, not in geographic distribution itself, a fact neither obvious nor widely appreciated: ‘I am perfectly aware that most of the secondary groups behave almost indistinguishably from primary ones in their fresh-water dispersal’ (in litt. to Darlington, 14 July 1948), yet, paradoxically, ‘it is… difference in [geographic] pattern…which has convinced me that the [six] groupings [primary, secondary, vicarious, complementary, diadromous, sporadic] have considerable basis in fact.’ (Myers 1949: 320). Madam How and Lady Why The coherence and influence of the New York School are too pervasive, throughout much of the 20th century, to remain unexplained or dismissed (Menard 1986: 84) even as ‘sophistry’ (Carey 1988: 133) or ‘solecisms’ (Carey 2000: 129). One suggestion, to have been ‘wrong for the right reasons’ (Laporte 1985, 2000), explains nothing about any mistake made in the interpretation of the biogeographic data,

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how ‘invalid’ inference derives from ‘valid’ principle (Laporte 1987: 331). Rather, the suggestion implies that there is no mistaken principle to understand and to explain— that to any question of earth history the biological data are either neutral (Oreskes 1999: 295) or they are incomplete and, therefore, unreliable (Frankel 1981, 1984; cf. Craw 1984c). If so, what then is the right reason, or valid principle, underlying Simpson’s (1966: 10) conclusion, based on data then judged reliable and complete enough, that for example ‘Australia has been in the fullest sense a dead-end’? There is no principle, none at all, beyond that of centre of origin. If ideas about centres of origin are method-generated artefacts of paralogous comparison, then the ideas and their mistaken basis, even Simpson’s, are to that degree rationally explained – the how but not the why (Kingsley 1870). Yet their artefactual nature was evident long before notions of geographic paralogy came into being (Cain 1944; Croizat et al. 1974; Nelson and Platnick 1984b). The only other explanation suggested (by Croizat 1981: 503) is that of Charles Darwin (1859: 352): ‘the simplicity of the view that each species was produced within a single region [centre of origin] captivates the mind’. So it did. And so it does still, until one learns better and moves on. Acknowledgments For comment, information and literature we are grateful to George Ball, Barbara Brown, Joel Cracraft, Chang Meemann, Daniel Cohen, Robin Craw, Jorge Crisci, Michael Crisp, Andrew Drinnan, Malte Ebach, Gregory Edgecomb, Lance Grande, John Grehan, Michael Heads, Wolfgang Hennig, Marianne Horak, Christopher Humphries, Philippe Janvier, David Johnson, Sylvia Kirchengast, Peter Linder, Jorge Llorente Bousquets, Malcolm McKenna, Stephen McLoughlin, Paul Manos, Bernard Michaux, Juan Morrone, Charles Myers, Lynne Parenti, Norman Platnick, Jay Savage, Scott Schaefer, Randall Schuh, Victor Springer, Dennis Stevenson, Jens Sommer-Knudsen, Pascal Tassy, François Vuilleumier, Jonathan Waters and David Williams. We are grateful also to William Cox and the Smithsonian Institution Archives for photocopies of correspondence between George Myers and W. D. Matthew, P. J. Darlington and W. K. Gregory. References Adams P (1999) ‘The stranger from Melbourne: Frank Hardy—a literary biography.’ (University of Western Australia Press: Nedlands) Anderson NM (1998) Marine water striders (Heteroptera, Gerromorpha) of the Indo-Pacific: cladistic biogeography and Cenozoic palaeogeography. In ‘Biogeography and geological evolution of SE Asia’. (Eds R Hall, JD Holloway) pp. 341–354. (Backhuys Publishers: Leiden) Andriashev AP (1987) Development of Berg’s concept of bipolarity of marine fauna. Biologiya Morya 2, 60–67. Anon (1957) K. P. Schmidt. Copeia 1957, 331.

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Manuscript received 11 April 2000, accepted 19 September 2000

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