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.ACorresponding author; email: [email protected]
Geography is impartial.Osmar White
Abstract. The many methods of biogeographic analysis proposed in recent years generate artefactual results thatimpede understanding, discovery and progress. Eliminating geographic paralogy from data reduces or eliminatesartefactual interpretation. Recent cladistic studies of extant Nothofagus agree in showing only three informativenodes relevant to intercontinental relationships. In cladistic representations of global distributions, Gondwana is ator near the base of the geographically informative nodes, which force Gondwana to appear as a centre of origin ofmodern life in general. Centres of origin are artefacts of comparison based on geographically uninformative andparalogous nodes. Postmodern revivals of dispersalism fail to acknowledge, explain, avoid, learn from and improveon the artefactual centres of origin of the 20th century dispersalism, as represented particularly by the New YorkSchool: 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).Vicari ance bi ogeography and the New York SchoolG. Nelson and P. Y. LadigesG. Nelson and P. Y. LadigesBT00025
Biogeography: a mess of methods
Biogeography has been reviewed, more or lesscomprehensively, several times in recent years. The reviewstestify to a continuing interest in their subject, stemming inthe 1960s from the revival of continental-drift theory andfrom the development of cladistic systematics. From thedecades since, the literature forms an interconnected whole,contemporary with the development, among other things, ofpostmodernism (Llorente Bousquets and Espinosa Organista1991; Crisci and Morrone 1992; Espinosa Organista andLlorente Bousquets 1993; Morrone and Carpenter 1994;Reynoso Rosales 1994; Enghoff 1995; Morrone and Crisci1995; Turner 1995; Morrone et al. 1996, 1997; Biondi 1998;Craw et al. 1999; Humphries and Parenti 1999; Vuilleumier1999; 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), therewas a symposium ‘Historical biogeography: a critique’. Thesymposium provoked the response that ‘Biogeography is amess’. As explained by its reporters, the meaning of theremark is that ‘biogeography is nowhere near finding aunifying and scientific method’ (Tassy and Deleporte 1999:14, translated). The meaning is similar to that ofobservations from earlier times (Keast 1977: 249) that ‘some
writers have recently been critical of the lack of a unifiedmethodology; e.g. Vuilleumier 1975’; also Rosen (1978) onVuilleumier (1978)—the centuries-old tension betweenecological and historical approaches to biogeography.
The varied possibilities to interpret data about taxa, theirinterrelationships and their geographic distributions, may beseen as different methods of analysis of data of these kinds(as in the summary diagram of Ebach and Edgecomb inpress: fig. 9). Different methods, even applied to the samedata, 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 aredriven to the view of Samuel Clemens (1897: 699) that ‘thevery ink with which all history is written is merely fluidprejudice’ (emphasis added).
Unavoidable it might seem, when faced with differentmethods, is choice among them. So, too, are hopes that onemethod might be better than others and that choice might beguided by objective criteria. If one method were preferred byall concerned persons, then there would be consistent resultsderived from the same data. If, however, its results wereartefactual to any degree, then a consistent method in generaluse would perpetuate the same problem, history as artefact ifnot prejudice, which would thereby have become obscured—swept under the rug of its apparent solution and the attendantforces of social conformity. Obscured, too, by the cloud of
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calumny likely raised around anyone using a methodconflicting with any currently in fashion. So testifies therecent history of cladistic systematics (Deleporte andLecointre 2000) and the emerging vanity that there be butone way, believed technically perfectible if not alreadyperfect enough, to skin the cat of systematic endeavour(Platnick et al. 1997).
On a broad scale, there is postmodern culture, itself aproduct of the 1960s according to one commentator,‘characterised by continual shifting surfaces and a new‘depthlessness’ derived from ‘the shock of slowly becomingaware 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 tofind a way to better understanding.
Geographic paralogy
In 1996 we suggested that artefactual results stem from datathat contain geographic paralogy (Nelson and Ladiges 1996).Use of the term ‘paralogy’ is by analogy with its use inmolecular systematics begun by Walter Fitch in the late1960s (Fitch 1970, 2000). There, paralogy refers tomisleading comparison between duplicated genes that havehad independent histories.
Geographic paralogy is evidenced for different taxa bytheir partly or wholly overlapping distributions—duplicatedgeographies of taxa that have had independent genetichistories. These matters reduce to two novel ideas: (1) that anode of a cladogram, representing a phylogeneticrelationship among organisms, is either geographicallyparalogous or it is not; (2) that any interpretation of aparalogous node is apt to be artefactual. We suggested thatgeographic data associated with non-paralogous nodes arethe only such data actually relevant to cladistic biogeography.In our experience, removing paralogy from data preventsartefactual interpretation, if not altogether then to asignificant degree. We expect that this will prove to be thecase also in the experience of other persons (e.g. Anderson1998) and time will tell.
Nothofagus—southern beech trees
For Darlington (1965: 140), Nothofagus was ‘the key to thehistory of terrestrial life in the far south’ because (p. 147) it‘is likely to disclose a geographic history that has beenfollowed by many other plants and by many invertebrateanimals’ (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 theCretaceous, crossed the tropics to Australia or New Zealandor both, radiated there and somehow made a triple dispersalhalf way around the southern end of the world’. This meansthree dispersals to explain the three subgroups recorded forSouth America (e.g. Van Steenis 1972: fig. 2; see alsoTakhtajan 1969: 154, 162; Whitmore 1981: 80; Hill 1992:
fig.1, 1994: fig. 16.1). The explanation implies that theSouth American representatives of each subgroup wouldprove related most closely to species of the jumping-offpoints for the three eastward dispersals. So far, this hasproved not to be the case and is contradicted by subsequentdiscovery, as if the three hypothetical dispersals really hadbeen from South America to the areas in the west (for ‘morethan three’ see Fleming 1963: 379; Cracraft 1975; Melville1982: 80). See for origin in North America and dispersal viaAsia and/or South America (Oliver 1925; Schuster 1976);
Fig. 1. (Left) Interrelationships of 30 extant species of Nothofagus,with their native area indicated: Aust, Australia; N Ca, NewCaledonia; N Gu, New Guinea; NZ, New Zealand; S Am, SouthAmerica, after Linder and Crisp (1995: fig. 3). (Right, above) Thethree geographically informative nodes (1–3) with relevant areas,derived from diagram at left. Nodes marked 0 are basal for paralogy-free subtrees (right). All other nodes are geographically paralogous.(Below) Combination of three informative nodes from above (Node 1represents 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 possibleresolutions. The tree is based on combined data: 26 morphologicalcharacters (Hill and Jordan 1993) and 46 informative characters fromrbcL sequences (Fig. 2). Analysed separately by Linder and Crisp(1995: fig. 1), the morphological data yield a strict consensus of12 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 fourtrees; of 19 nodes, three are geographically informative, equivalent tothe combination (right, below) with S Am sister to NZ-Aust. Also onthe basis of morphology, an earlier ‘tentative phylogeny’ shows twogeographically informative nodes in one subtree: ((N Ca N Gu) NZ)SAm (Melville 1973: fig. 5, Cracraft 1975: fig. 5, Humphries 1981b:fig. 21.7). One of Humphries’ trees (fig. 21.9) has one geographicallyinformative node: (N Ca N Gu) NZ Aust S Am and the other tree(fig. 21.10A) has none at all.
Vicariance biogeography and the New York School 391
Eurasia and dispersal via Africa–India (Raven and Axelrod1972, 1974: 573); ‘an area of Gondwanaland which includedChile and Patagonia, Western Antarctica and the NewZealand platform’ (Melville 1973: 444); ‘a region betweenNew Zealand, Antarctica and Australia’ (Hanks andFairbrothers 1976: 69); Antarctica (Moore 1972; Dettman1989, 1994; Dettman and Jarzen 1990; Dettman et al. 1990);‘the region encompassed by southern South America and theAntarctic peninsula’ (Hill 1996: 247, Hill and Dettman1996: 13; Hill et al. 1996: 182, 1999: 283; Veblen et al. 1996:
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 thestrict consensus tree of Linder and Crisp (1995: fig. 3). Of 27nodes in the tree, based on morphology and molecularsequences, only three are geographically informative(Fig. 1). Nodes 1 and 2 relate Australia and New Zealandmore closely than to South America. Node 3 similarly relatesNew Caledonia and New Guinea (Ladiges et al. 1997: 128;Ladiges 1998: 236–237). Nothofagus is not richlyinformative about intercontinental relationships. The threenodes, nevertheless, are mutually consistent even if
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 isshown in place of N. codonandra and N. discoidea). (Right) The threegeographically informative nodes (1–3) with relevant areas, derivedfrom diagram at left. The tree is a strict consensus of 53 trees(according to Manos 1997: 1142), based on 46 informative charactersfrom 1345 bp of chloroplast DNA rbcL. Linder and Crisp (1995:fig. 2) find a different strict consensus of 54 trees. Of 15 nodes, twoare geographically informative (equivalent to Nodes 1 and 2).
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) withrelevant areas, derived from diagram at left. The tree is stated as astrict consensus of six trees, but shows only one collapsed node, witha maximum of three possible resolutions (one shown by Manos 1997:fig. 6; Swenson et al. 2000b: fig. 2, which includes two other NewCaledonian species in place of N. aequilateralis). The tree is based oncombined data: 1345 bp of rbcL (Fig. 2), 588 bp of ITS nrDNA and 23morphological characters (modified from Hill and Jordan 1993).Analysed separately, the morphological data yield a strict consensus ofeight trees (Manos 1997: fig. 3). Of 13 nodes, two are geographicallyinformative (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) moreclosely than to subtree 1. Node 7, leading to the basal nodes ofSubtrees 2 and 3, is paralogous and geographically uninformative.
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 astrict consensus of 31 trees, based on 146 bp of rbcL and 453 bp ofatpB-rbcL intergenic spacer.
392 G. Nelson and P. Y. Ladiges
minimally corroborative: one node’s worth of corroboration.These results are similar to those, paralogy aside, ofHumphries et al. (1986: fig. 4.14; cf. Patterson 1981 andHumphries 1981a, 1981b, 1983 for both of whom‘Nothofagus is uninformative on the interrelationships ofSouthern Hemisphere areas’).
The molecular sequences are for 23 of the 30 species, forwhich Martin and Dowd (1993: fig. 2) published a strictconsensus. Of 16 nodes in the tree, only three aregeographically informative (Fig. 2). The three nodes have thesame 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 newmolecular sequences. Of 15 nodes in the tree, only three aregeographically informative (Fig. 3). The three nodes have thesame 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 molecularsequences, some new. For the combined data, a strictconsensus of 19 nodes (1997: fig. 5) has only threegeographically informative nodes (Fig. 4). The three nodeshave the same information as the three nodes of Linder andCrisp. For the new molecular sequences analysed separately,Manos (1997: fig. 2) published a strict consensus. Of19 nodes in the tree, only three are geographicallyinformative (Fig. 5). Node 2 differs from those above inrelating Australia and South America more closely than toNew Zealand.
For a similar combined analysis of the 22 species, Jordanand Hill (1999: fig. 2) published a strict consensus(19 nodes), having three geographically informative nodeswith the same information (Fig. 6). The addition of six fossilspecies from Tasmania yielded a strict consensus (1999:fig. 4, 17 nodes) having only two geographically informativenodes (Fig. 7), with the same information (Australia andNew Zealand related more closely than to South America).The third node above, relating New Guinea and New
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 strictconsensus of two trees, based on 588 bp of ITS nrDNA.
Fig. 7. (Left) Interrelationships of 22 extant, and from Tasmania (T)six fossil, species of Nothofagus, with their native area indicated (seeFig. 1), after Jordan and Hill (1999: fig. 4). (Right) The twogeographically informative nodes (1–2) with relevant areas, derivedfrom diagram at left. The tree is a strict consensus of 90 trees, basedon combined data (Fig. 6).
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 astrict consensus of two trees, based on combined data: 24 morpho-logical characters and 121 informative characters from sequences ofrbcL (Fig. 2) and ITS (Fig. 5).
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Caledonia more closely than to South America, is renderedparalogous by the placement of two fossil species fromTasmania (N. mucronata, N. lobata), the former within aNew Guinea–New Caledonia clade in an unresolvedpolytomy with six extant species, the latter within a resolvedSouth America clade as sister to one of four extant species(N. nitida).
Paralogy v. local precision
One cause of paralogy is ‘imprecise characterisation ofgeographic areas’ (Nelson and Ladiges 1996: 11–12). Areascommonly used to describe intercontinental relationships ofNothofagus (South America, etc.) are not precise. Species ofNothofagus have more or less discrete distributions withineach such area. More precise characterisation woulddoubtless expose certain nodes as non-paralogous,interrelating local areas within South America, NewZealand, Australia, New Guinea and even New Caledonia.
Paralogy, corroboration, consistency and conflict
By logical necessity, different subtrees imply (interconnectby) paralogous nodes. If informative of geographicrelationship, different subtrees are corroborative, consistentor conflicting. Corroboration occurs when different subtreesinclude the same three (or more) areas interrelated in thesame way by one (or more) non-paralogous node. Conflictoccurs when such areas are interrelated in different ways.Consistency with no corroboration occurs when subtreesinclude fewer than three such areas and logically combine ina single and more informative tree. Without paralogy (in ageneral sense inclusive of molecular systematics), there is nopossibility either of corroboration (Nelson 1994:138) or ofconflict.
Nothofagus, ratite birds and Gondwana
Craw, Grehan and Heads recently contrasted Nothofagus andratite birds – kiwis and their relatives (1999: 27; also Craw1983, 1985; Heads 1985, Grehan 1988b; Page 1989):
Distribution patterns of the ratite birds and thesouthern beeches are in no way biogeographicallyhomologous or congruent, nor were they oncemembers of a widespread, ancestral Gondwana biotaas is often suggested in the biogeographical literature(e.g. Humphries 1981a; Patterson 1981). Nothofagusis a member of a non-Gondwanan trans-Pacificfagalean alliance; the ratites are a Gondwanic groupcentered on the South Atlantic and Indian Oceanbasins. In the Southern Hemisphere these groups aregeographically sympatric only in southwestern SouthAmerica and in eastern Australasia where their tracksintersect.In their view, Gondwana is evidently understood to
embrace taxa with distributions across the present Atlanticand Indian Oceans but not across the Pacific, as if all
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; Grehan1988a: fig. 3; Henderson 1989: fig. 15; Llorente Bousquets1991; 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 andGibson 1983: fig. 9.8; Crisci and Morrone 1989: fig. 4;Grehan 1991: fig. 8, 1993: Fig. 3, 1995: fig. 7; Brown andLomolino 1998: fig. 12.3; Cox 1998: fig. 2; Craw et al. 1999:figs 6–12, 7–2; Morrone 2000: fig. 3). At the most generallevel, its cladistic representation shows five nodes: Atlantic,Indian, Pacific, boreal and austral with no relationshipsshown among the five (Nelson 1985: fig. 1; Chow et al.1996: 267). The last two, boreal and austral, have long beenassociated as bipolarity (Berg 1933; Du Rietz 1940;Andriashev 1987), which would form an additional, sixth,node to the cladistic representation (Fig. 8A, B). There isreason to associate bipolarity with the Pacific rather thanwith the Atlantic and Indian Oceans and this associationwould form an additional, seventh, node (Fig. 8A, B), the‘Pacific Rim’ of Craw et al. (1999: 27). Bipolarity, evenextended to the Pacific Rim and beyond to the Caribbean(Rosen 1976; cf. Briggs 1994), does not exhaust thecomplexity 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 Atlanticand 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, Gondwanaaccording to Craw et al. (1999), with alternative placement shownin B.
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the meaning of the term ‘Gondwana’. With respect to ratitesand Nothofagus, their evidence is non-existent. Ratites mightbe Atlantic and Indian and Nothofagus Pacific. It does notfollow that the two (Atlantic, Indian) relate more closely thanto the one (Pacific). Craw et al. note (p. 26) that ‘Nothofagusis 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 ofGondwana, but they amply drape over her left arm andshoulder.
Bony-tongue fishes and Gondwana
We refer here to South America and two extant freshwaterfishes of the bony-tongue family, Osteoglossidae: thearawana (Osteoglossum) and the pirarucú (Arapaima) ofAmazonian white water (Fig. 9). They live together over avast distribution (Berra 1981). It is said that the smallerarawana is the ‘favourite bait’ of the larger pirarucú (Herald1961) or is sometimes eaten (Roberts 1972), but is notalways recorded as an item of the natural diet (Lüling 1964,1971). The pirarucú, one of the world’s giant fishes, has itsnearest living relative in Africa (Heterotis, tropical WestAfrica and the upper Nile; Roberts 1975). The arawana hasits nearest relatives in a complex of species (Scleropages):one or more species in Sumatra, Borneo and South East Asiafrom Malaya into Thailand; two or more species in NewGuinea and tropical Australia, which have the name saratogain recent Australian books (Merrick and Schmida 1984;Grant 1987; Larson and Martin 1990) and burramundi orbarramundi in earlier books (Whitley 1960; Munro 1967;Lake 1971, 1978; Pollard 1980), a name now reserved incommerce for an unrelated fish (Anon. 1995; Yearsley et al.1999). The New Guinea and Australian forms are relatedmore closely than to the Malayan species (on the basis ofpersonal observations). Possibly they are more closelyrelated to the South American arawana than to the Malayanspecies (Nelson 1969a). More remote are relatives, fossil inIndia (Bonde 1996) and extant and fossil in Africa (Li andWilson 1996: fig. 4).
The relevance of the fishes is the evident trans-Pacificrelationship within a group, evidently Gondwanan, withfossil roots into the mid-Mesozoic (Kumazawa and Yoshida2000). In terms of their geography, the fishes are nearlyequivalent to the ratites and Nothofagus combined, with atrans-Pacific distribution extending beyond the shoulder intoIndia and Africa, the heart of Gondwana. So in SouthAmerica is the pirarucú a Gondwanan element and thearawana not, because the one relates to the east and the otherto the west?
Trans-Pacific relationships, Gondwana and the world
Craw et al. (1999: 154–160) consider Pacific distributionand Croizat’s (1960: fig. 8) concept of a geologicallycomposite New World (reproduced in Craw 1984a: fig. 2,
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). InCroizat’s view, the western sector, from Alaska to Chile,relates to lands bordering the West Pacific, but not in theirview to Gondwana. Yet relevant lands of the West Pacific areseen as physical parts of a geologic Gondwana (Metcalfe1999; Metcalfe et al. 1999; Michaux and White 1999). So,too, are Pacific terranes—harbouring trans-Pacificbiological relationships—seen as physical parts of a geologicPacifica continent and of a more inclusive Proto-Gondwanalandmass (Craw 1985: fig. 5, 1988: fig. 13.9, 1991: fig. 5;Chow et al. 1996: 253).
Are trans-Pacific biological relationships external to orembedded within Gondwana and her history? As abiogeographic relationship of today, is Gondwana to beunderstood to relate distributions across the Atlantic andthose across the Indian Oceans? Or, as an alternative andinformative Node 8, to relate distributions over one or theother possible combination of oceans, Indo-Pacific orAtlanto-Pacific? In Croizat’s summary, the only other nodeavailable for the name ‘Gondwana’ is the cladisticallyuninformative basal node of life’s global distribution(Fig. 8B). In that sense Gondwana and today’s world wouldbe one. This alternative implies that the life of the pastGondwana is reflected today in the distribution of life theworld over, a fair summary of the actuality.
For a modern distribution to appear Gondwanan, ratherthan merely Atlantic, Indian or Pacific, there seem to be tworequirements: that it involve more than one ocean basin; andthat in a phylogenetic tree it associate with a node, likely to begeographically paralogous, that is basal enough so that allmore basal nodes are also paralogous. Such nodes also inviteassociation with Gondwana. These requirements (and nodes)force Gondwana to appear, artefactually, as an ultimate centreof origin of modern life in general (e.g. Cracraft 2001).
Fig. 9. Interrelationships and distribution of extant fishes of thefamily Osteoglossidae, including three species of Scleropages, withtheir native area indicated (see Fig. 1 and text). Nodes: 0,Paralogous; 1, Atlantic; 2, Gondwana; 3, Pacific; 4, Arafura Sea—Torres Strait.
Vicariance biogeography and the New York School 395
Centre of origin?
Faced with paralogy, various notions self-destruct. Onenotion, perhaps the most notable casualty, is again, the centreof origin, which in a cladistic context seems always a productof paralogous comparison (Ebach 1999).
Consider a hypothetical example (Fig. 10), sent to WilliHennig a few years before he died in 1976. There are twotaxa (extant or fossil) in South America, one related to a thirdin Africa (Nelson 1974: fig. 2). The question posed toHennig was: is there evidence, however slight, of a centre oforigin 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; Brundin1966: 56)—the usual cladistic criterion (Fig. 11; Ross 1974:214–220; Platnick 1981; Wiley 1981; Nelson and Platnick1984a); no, if the basal node be set aside as paralogous.
Is it reasonable in this hypothetical case, despite theduplicated geography in South America, to consider that theancestor of the three taxa occurred also in Africa? Yes, whynot? The cladistic criterion used to resolve a centre of originis minimality of implied dispersal events (Nelson 1969b).Why not minimise them to zero?
Without a centre of origin and dispersal therefrom, whatis the implied history in this hypothetical case? That the firstsplit of the widespread ancestor isolated a population inSouth America before the later split between the lesswidespread South American–African population.
Paralogous nodes—geographically informative?
A second notion is that attached to each node of a cladogramthere are relevant geographic data deserving of interpretation—not so for paralogous nodes. There is no necessity to
interpret geography to fit paralogous nodes in the hope thata meaningful pattern might emerge from their analysis evenby a computer program.
Geographic homoplasy?
A third notion is that apparently conflicting and artefactualgeographic data are analogous to homoplasy in systematicdata, which, hopefully, is distributed randomly, more or less,among the nodes of a cladogram—not so becausegeographically paralogous nodes increase towards the baseof cladograms generally.
Consider two nodes: one node (non-paralogous) relatinga 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 insectsupon which they feed. The former is geographically relevantand the latter not. The former node speaks to a relationshipbetween the river basins. The latter to who knows what?
Creation myth: past and present
A remarkably recurrent theme of 20th century biogeographyis the quest for origins of taxa and of their distributions, or inother words ‘The geographic origins of individual taxa anddispersal 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 creationmyth of the Bible – the Garden of Eden. Linnaeus and laterwriters used the myth as an explanatory device ofbiogeography (Nelson and Platnick 1981; Nelson 1983). Itpersists in the ‘out of Africa’ sagas of human origins andtheir molecular embellishments. It is currently underpostmodern revival: if the flora of New Zealand, ‘isolated
Fig. 10. In South America, two taxa (extant or fossil) of which onerelates 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 whichone relates to a third in Africa, illustrating minimum evidence(paralogous Node 1) to resolve a centre of origin according to theusual cladistic criterion (Hennig’s Progression Rule). (Middle) Twicethe minimum evidence (paralogous nodes 1–2). (Bottom) Thrice theminimum evidence (paralogous nodes 1–3). Simpson (1947: 649)regarded such evidence (‘a variety of earlier relations’) as ‘morereliable than…earlier appearance’ in the fossil record.
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from Australia and Antarctica...for 60 million years...can beshown to have arrived over the sea...then biogeographichypotheses the world over which involve any kind of landconnection must be reconsidered’ (Pole 1994: 625). Theargument is the familiar ‘if-then’ of the ‘professionalbiogeographer’ (Darlington 1964a: 1084): ‘if Nothofagushas been wind-dispersed across southern ocean gaps in thelate Cretaceous and Tertiary, [then] Glossopteris may havebeen 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’, bio-geography is now seen suddenly to have ‘got real’ (Heaney1999: 435–436):
For me, for example, knowing with good confidencethat the continental rocks included in the Philippineisland of Mindoro were covered by marine seas at thetime of their fragmentation from the Asian continentand that they did not re-emerge as part of a subaerialisland until the late Miocene (roughly 10 million yearsago) makes all the difference in successfully sortingprocess out of patterns of distributions.The possibility of complete submergence of New Zealand
in the late Oligocene and of New Caledonia in the earlyTertiary, is similarly used to argue for ‘entirely long-distancedispersal’ as explanation of the biota of these islands (Pole1994: 628–629). Once again (Darwin 1845: 378) ‘we seemto be brought somewhat near that great fact—that mystery ofmysteries’…the vision that ‘the floras and faunas of manyislands, including New Zealand...must have crossed watergaps’ (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-distancedispersal (McGlone et al. 1996: 83)—the ‘flotsam of sea andsky’ (Cranwell 1963: 387):
...obvious candidates are storm-force winds and thefeet and plumage of birds. While the probability of thishappening in any one year is likely to be extremelysmall, migrational lags of millions of years may haveprovided the requisite time interval for the improbableto happen.
(cf. Matthew 1915: 206–209, 1939: 37–40; Simpson 1952;Darlington 1957: 484).
Emerging lifeless from the ocean, barren lands feature inLinnaeus’ explanation of life’s distribution throughout theworld (Linnaeus 1744, 1781; Browne 1983; Frängsmyr1983; Seberg 1985; Larson 1986, 1994; Rupke 1996, 1997;Papavero et al. 1997; Bueno et al. 1999; Llorente Bousquetset al. 2000; Bueno Hernández and Llorente Bousquets2001). Whether universally or locally applied, the vision andits effect are the same—to render superfluous any geogra-phic comparison across taxa. Biogeography thereby reducesto dispersalism and its empirical impossibilities—marking
the centre of origin and time of arrival of each, or even one,propagule theorised to have been successful in colonising thebarren landscape to the east, or to the west, of Eden. To allappearances, a real problem (what to believe? or how toproceed? or when and where to stop?) is thereby solved. Thatthe history of biogeography is littered with solutions of thekind testifies to their continuing popularity (Nelson 1978).
Popular, too (Macphail 1997b: 425), is the modernresonance with the classical dictum, from Aristotle (HistoriaAnimalium, book viii, s28) via Pliny (Naturalis Historia,book viii, s17), that ‘out of Africa there is always somethingnew’—Ex Africa semper aliquid novi (Smith 1939). Themodern resonance stems from Darwin’s (1871: 199)comments on the ‘Birthplace and Antiquity of Man’:
It is therefore probable that Africa was formerlyinhabited by extinct apes closely allied to the gorillaand chimpanzee; and as these two species are nowman’s nearest allies, it is somewhat more probable thatour early progenitors lived on the African continentthan elsewhere.The logical principle of Darwin’s ‘cogent reasoning’
(Leakey 1960: 18) later became generalised as theprogression rule to circumscribe a centre of origin fromphylogenetic trees, including those based on fossil as well asmolecular data (Stoneking 1996: fig. 11.4; Patterson 1999:fig. 16.1). The principle underpins the answer to thequestion, ‘Where are we to pitch the centre of dispersal? Theevidence, as it stands today, favours Africa…If we may selectone region as more likely than another, then our choice fallson the uplands of Uganda and Kenya’ (Keith 1948: 214). Theevidence is paralogous nodes, or the equivalent nodes ofpresumably direct ancestry (Nelson 1973: fig. 1).
Recast by Louis Leakey as ‘Charles Darwin’s prophecy’(also Tobias 1984: 37), the modern resonance peaked in aseries of fossil finds (Tattersall 1995), beginning mostnotably in 1924 with (Leakey 1974: 193) ‘Raymond Dart’sspectacular discovery of the juvenile skull of Austra-lopithecus at Taung, South Africa’:
I stood in the shade holding the [fossil] as greedily asany miser hugs his gold, my mind racing ahead. Here,I was certain, was one of the most significant findsever made in the history of anthropology. Darwin’slargely discredited theory that man’s early progenitorsprobably lived in Africa came back to me. Was I to bethe 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 butindisputable outline of the human emergence on the Africanhighland’ (Ardrey 1961: 27). So developed the story of ‘thatgreat fact—that mystery of mysteries—the first appearanceof new beings on this earth’ (Darwin 1845: 378).
Associated with Louis Leakey (1903–1972) is anotherprophecy, which he did not survive to judge at the stated time
Vicariance biogeography and the New York School 397
—that the future, even 10 years hence, would see ‘how littlewe knew, how stupid we were in 1965!’. His doubt extendedeven to the ideas that the human family ‘Hominoidea startedin Africa’ and, accordingly, that ‘there were…movementsout into Europe and Asia’ (Leakey 1965: 17, 1972: 399). By1969, however, Leakey’s doubt was seemingly forgotten(Leakey and Goodall 1969: 170):
Charles Darwin’s prophecy is coming true. More andmore evidence is accumulating which points to theAfrican continent and particularly the East/CentralAfrican 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 McKie1996), the ‘footsteps of Eve’ (Berger 2000), the ‘Genesischronicles’ (McBride 2000). The question naturally arises(Foucault 1977: 144): ‘does this not form a history, thehistory of an error that we call truth?’.
New York School
Of dispersalism, its history in the century past belongs alsoto what Croizat (1958: xi) called the ‘New York School ofZoogeography’, in reaction to which vicariance bio-geography developed (Nelson and Rosen 1981). AmongNew Yorkers, the principal players in the ‘great and splendiddrama’ are fossil vertebrates, mammals in this case andpersons posing as their interpreters: primarily William DillerMatthew (1928: 54; Bowler 1996) and secondarily GeorgeGaylord Simpson. In their view the crucial evidence isprovided by discovery of the fossil ancestors of a taxon—plesiomorphic fossils. Once found, fossil ancestors—the‘true genetic sequences’ (Matthew 1925: 288)—revealdirectly the centre or place of origin of the taxon in question:the geographic space inhabited by the ancestors. That thisempirical discovery was possible, even ‘demonstrably true’(Simpson 1937: 253), they took for granted and neverquestioned. When occasionally this empirical discoveryseemed not forthcoming, they ‘somewhat sadly’ lamented itsabsence (Simpson 1978b: 324). It was early questionedduring the development of cladistics and found impossible toachieve. Years of argument and discussion of the matter werelater summarised by Colin Patterson (Fortey 1999; Foreyet al. 2000) with his usual clarity and eloquence: ‘Fossilsmay tell us many things but one thing they can never discloseis whether they were ancestors of anything else’ (Patterson1978: 133, 1999: 109).
W. D. Matthew (1871–1930)
Of Matthew it is said that ‘the stray lock of hair and thealmost inevitable cigar were trademarks of the man’ (Colbert1992: fig. 40). So wrote Ned Colbert, who in 1933 marriedWilliam Diller’s younger daughter Margaret. Matthew wrotea celebrated essay of 150 pages, ‘Climate and Evolution’.Thirty years later, Karl Schmidt (1943: 242) wrote that the
essay had inaugurated ‘a new phase in the study of animalgeography’. Thirty years later still, Simpson (1978a: 272)described it as ‘one of the most seminal or heuristic studiesof paleogeography and historical biogeography’. The essaywas published in 1915 by the New York Academy ofSciences (Matthew 1915). Nearly 25 years later it wasreprinted in 1939 (Matthew 1939) because, according toColbert (1939: v) at the time, ‘there has been a steadydemand for this paper that has continued to the present day,long after the original edition had been exhausted’. The essaywas 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 dispersalsouthward: to Africa, through South East Asia to Australiaand across a Bering land bridge to and through NorthAmerica to South America. His is the same notion thatoriginated in the mid-19th century if not earlier. ErnstHaeckel (1834–1919) used it to explain the history ofhumans and their wanderings from their centre of origin inParadise (Haeckel 1889: pl. 20 and later editions, modifiedfrom Haeckel 1879: pl. 15 and earlier editions beginningwith the second [the many English translations begin withHaeckel 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 havedisagreed (also Savage 1958: 154), pointing out correctlythat his centres of origin are north of the languid tropics. Inhis centres, ‘greater activity and higher development of life’would emerge in reaction to ‘the inclemency of nature, thescarcity of food, the variations of temperature, as well asagainst 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 outlinedfirst by Buffon [1707–1788] and was elaborated by [AlfredRussel] Wallace [1823–1913] in his great book on [TheGeographical] Distribution of Animals [1876]’ (noted alsoin Croizat 1958: i).
K. P. Schmidt (1890–1957)
Matthew was seen as an inspiring adversary of continentaldrift. In the words of one of his students:
In the very year when [Alfred] Wegener proposed histheory [Wegener 1915], this...was shown by WilliamDiller 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 thethumb by a small snake brought to the Museum foridentification from the local zoo (Anon. 1957; Davis 1959).He wrote also (pp. 780–781):
A group of students in zoology and paleontology underWilliam King Gregory at the American Museum of
398 G. Nelson and P. Y. Ladiges
Natural History in the years 1915 to 1927 came alsounder the influence of W. D. Matthew...The groupincluded Alfred Sherwood Romer, Charles LewisCamp and Gladwyn Kingsley Noble, with many othersand a little more indirectly Emmett Reid Dunn andmyself. All of us have remained disciples ofMatthew...we have tended to look a bit askance at those‘who knew not Matthew’; and it had not escaped someof our colleagues that Matthew’s work had become akind of Holy Writ to his disciples...The leadershipamong the group of Matthewsians has now somewhatnaturally fallen to George Gaylord Simpson, whosucceeded Matthew in the position of Curator incharge of Vertebrate Paleontology at the AmericanMuseum in 1944.
G. G. Simpson (1902–1984)
In 1924 while a graduate student at Yale University, theyoung Simpson was hired as assistant to Matthew, spending6 weeks with him in field work in Texas (Laporte 1986,1990). In 1927 Matthew left New York City for theUniversity of California at Berkeley, there to found ‘the onlyseparate department of paleontology at an Americanuniversity’ (Colbert 1989: 143). Simpson replaced him inemployment at the American Museum of Natural History(on Matthew’s recommendation) and in 1945 at ColumbiaUniversity (Hecht et al. 1972; Laporte 1991). Simpsonremarked: ‘I came to the American Museum to follow, at agreat distance, in the footsteps of W. D. Matthew’ (Simpson1945: vii). ‘He was a great paleomammalogist, a hero to meand with him familiarity bred only respect and admiration’(Simpson 1984: xvi). ‘I absorbed every word he spoke andread...every word he had written’ (Rainger 1991: 213).‘I soon was drawn to the essentials of W. D. Matthew’sviews’ (Simpson 1976: 8). And with Simpson, as early as1940, it was more of the same: ‘the general type ofgeographic history assumed by Matthew to be typical formammals is...here more explicitly supported’ (Simpson1940a: 141). ‘Matthew’s theory does afford the bestexplanation so far proposed’ (Simpson 1940b: 765). ‘Thedistribution of mammals definitely supports the hypothesisthat continents were essentially stable throughout the wholetime involved in mammalian history’ (Simpson 1943: 29).
Simpson, too, eventually wrote an essay, of a modest andless celebrated 64 pages, ‘Evolution and Geography’, firstpublished in 1953, reprinted a half-dozen times by 1968 andalso translated into Spanish (Simpson 1964) and French(Simpson 1969). It concludes: ‘All the biogeographicfeatures in the known history of mammals are best accountedfor on the theory that the continents have had their presentidentities and positions...the conclusion seems to apply notonly to the biogeography of mammals but also to that of allcontemporaneous forms of life’ (Simpson 1953: 61–63,1964: 55, 56, 1965: 130, 1969: 106, 107).
After moving from the American Museum to the Museumof Comparative Zoology (MCZ) at Harvard University,Simpson remarked (1961: 1684): ‘Matthew...has successorswho have followed in his footsteps and have, with constantlyimproved data, gone well beyond him. The main outlines...ofmammalian faunal evolution are now well established....Currently accepted general principles of historicalbiogeography...are derived largely from paleomammalogy inthe tradition of Darwin and Matthew. This is evident, forinstance, in a fine recent treatise...written by an [MCZ]entomologist, [Philip Jackson] Darlington [1957].’ Twentyyears later Simpson (1980: 253) described this as an‘excellent...treatise...on biogeography that is now out of datebut better than most of the recent works on that subject’—thereview by Keast (1977) being ‘incomparably the best modernsummary of the whole subject’ (Simpson 1978c: 219).
P. J. Darlington Jr (1904–1983)
Other forms of life is the theme previously taken up byDarlington in papers featured in the Quarterly Review ofBiology in 1938 and 1948. The former considers dispersal oforganisms to the Caribbean islands by means of wind,cyclonic storms (hurricanes), waterspouts and oceancurrents, and finds (p. 197):
The fauna is…very orderly…[The] animals are stilldistributed along the migration routes by which theirancestors reached the islands and this is taken to showthat the fauna is an accumulation of immigrantsderived from the mainland across water…No otherhypothesis will fit the facts.For George Myers (1954a: 19) the former was ‘an
extremely important paper on the origin of the GreaterAntillean fauna, in which [Darlington] concluded that [that]fauna gives no indication of a continental connection’. Thelatter paper, ‘The Geographical Distribution of Cold-Blooded Vertebrates’, prompted Karl Schmidt to remark(1955: 780):
Fortunately we have now had a strong cross-lightthrown on the main thesis of Climate and Evolution bya nondisciple [of Matthew], P. J. Darlington, Jr, of theMuseum of Comparative Zoology at HarvardUniversity, who reviews the whole matter from theevidence of the freshwater fishes, amphibians andreptiles.Darlington’s approach was later perfected in another
celebrated work. Considered ‘the most meritorious work inzoology published during the year’, the book‘Zoogeography’ (Darlington 1957) was awarded the 39thDaniel Giraud Elliot Medal of the National Academy ofSciences of the United States (NAS Website 2000; Carpenter1985: 10, but in the year 1957 not 1969). Not merely forcold-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
Vicariance biogeography and the New York School 399
New York School near the end of its development. The bookmarked the ‘end of an era’ (Savage 1958: 155), but not theend of that school’s influence, which, ‘in a virtuallyunassailable position’ (Simpson 1958: 379), continues to thepresent, ‘hardened into solid theory’ (Brown 1985: 15), mostnotably 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 historicalarchives’ [Darlington 1980: 11], 1964b: 970):
Darwin considered the evidence he had and decidedthat as far back as he could see the main pattern of landhad been the same as now...Fifty-six years laterMatthew, with much more evidence, reached the sameconclusion, but saw farther back and in much moredetail than Darwin could. And now, with still moreevidence, we can see still farther back and in still moredetail than Matthew could, but the conclusion is stillthe same.The existing pattern has evidently been formed by verycomplex movements of animals over the worldapproximately as it is now, not over extraordinary landbridges or drifting continents.
G. S. Myers (1905–1985)
Central to the ichthyological history, an academic conduitfrom New York to Cambridge and to the world at large—even to New Zealand (McDowall 1988: vii)—is GeorgeSprague Myers and his ‘more than 104 graduate and specialstudents’ at Stanford University, during his 35-year tenure asProfessor in the Department of Biological Sciences (1936–1970, Herald 1970). Earlier (1922–1924), Myers had been avolunteer assistant at the American Museum of NaturalHistory. It is said (Walford 1970: 3, Cox 1988: 3) that hethere came under the influence, among other persons, of KarlSchmidt, himself employed intermittently at that institution,eventually resigning as Assistant Curator in 1922 ‘to accepta position in Chicago as the first head of a new Division ofAmphibians and Reptiles at the Field Museum of NaturalHistory’ (Myers 2000).
Before Matthew’s death in 1930, Myers (1938: 341) hadapparently ‘gathered the ichthyological information’relevant to ‘the proposed enlarged and revised [second]edition of ‘Climate and Evolution’.’ When that secondedition was published (Matthew 1939), it was without hiscontribution, which had appeared shortly before (Myers1938). Therein he divided true freshwater fishes intoprimary and secondary classes. More numerous in species,the primary freshwater fishes ‘possess an ancientphysiological inability to survive in salt sea water, whichbinds them to the land as securely as any known animals’.Most primary fishes are in the group Ostariophysi, whichinclude carps, characins, catfishes and electric eels.
Secondary freshwater fishes ‘are generally restricted to freshwater but occasionally enter the sea’ (Myers 1938: 342–344).Other ‘still more salt-tolerant’ fishes found in freshwater helater grouped into four additonal classes (Myers 1949,1954a, 1954b; Kessel 1970 explains dates of Myers’publications). He stated that his accounts of the primaryfishes (1938: 354):
...make it clear that our present knowledge of the fishesvery distinctly favors a late Mesozoic or very earlyTertiary South Atlantic land connection and makes adirect northern origin...seem exceedingly unlikely. ButI refuse to take a definite stand on these questions. Itwould be extremely presumptuous, on the basis of thefishes alone, to attempt a flat contradiction of theHolarctic dispersal of the mammals, reptiles andamphibians 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 onnearly all recent American zoogeographers and but little onanybody else’. By 1946, ‘after eight years’ cogitation onfresh-water fish dispersal’ Myers had ‘gone over to thenorthern origin idea, with all the speed and grace of a catdragged backward by the tail over the dining room rug. Thereis simply no other answer to the fish distribution problem andI say this with perhaps fuller knowledge of the objectionsthan anyone’ (Myers in litt. to Darlington, 12 November1946). In 1949, he considered Matthew’s 1915 essay as ‘themost important single contribution to zoogeography sincethe time of Wallace’ (Myers 1954a: 19), a judgmentsurpassed only by Darlington (1959b: 488): after ‘the twochapters on geographical distribution and…parts of otherchapters’ in the ‘Origin of Species’ (1859) ‘the next reallyimportant 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 thesedivisions in an extensively well-thought-out general schemeof fresh-water fish distribution’. He noted that ‘in 1948,Dr Darlington gave an excellent resumé of the world’s fresh-water fish distribution, in which he expanded the fish datagiven in my 1938 paper and included amphibians andreptiles’ (Myers 1951: 11, 1954b: 39). For his partDarlington (1948: 4), as he had done 10 years before (1938:291), acknowledged that ‘I am greatly indebted to Dr GeorgeS. Myers for guidance in my work on fishes...’. Of hisdivisions Myers wrote to Darlington that ‘I have made onediscovery worth general quoting in my life, which I amhuman enough to want to see recognised. You did.’ (in litt. toDarlington, 17 April 1951).
Of Myers’ 1938 paper it was noted (Walford 1970: 12)that ‘20 years after publication this paper was acknowledgedby P. J. Darlington in his great book ‘Zoogeography’ as theprime reference on which he built that part of his book
400 G. Nelson and P. Y. Ladiges
dealing with fishes’. For his part Darlington (1957: x–xi)conceded that ‘Thanks to hard work on my part andespecially to aid from George S. Myers and otherichthyologists, what I shall have to say about fresh-waterfishes 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 notionsof continental stability in favour of drift as explanation ofgeographic distribution of fishes: ‘I believe continentaldrift 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 bepregnant, but only a little bit (1964a: 1090, 1979: 345):
Africa and South America probably were unitedbut...separated not later than the Triassic and perhapsearlier, so long ago that no clear traces of the union arevisible in distribution of existing life. Other southerncontinents were probably not united.Myers rejected continental stability because, with
increased knowledge of Central American fishes (orincreased meditation upon their significance), he againfound implausible the Matthew–Darlington explanation ofthe primary freshwater fishes of South America – theirdispersal through North and Central America from an Asiancentre of origin. Ironically, this rejected explanation, so clearand comprehensive, was due partly to Myers’ own prioreffort and history of shifting conviction (Walford 1970: 12).It is said (Géry 1969: 833) that ‘Darlington’s (1957)hypothesis...expressed rather faithfully the views ofspecialists with G. S. Myers at the head...’. And only 10 yearsbefore according to Schmidt (1955: 780):
The very important evidence of the freshwater fishes,long regarded as a proof of the necessity for past directconnection of Africa and South America, is nowregarded by George S. Myers as explainable by long-term round-about emigration via the Bering Bridge[and Central America] rather than by hypotheticaltrans-Atlantic connections.In the 1960s Myers did not adopt a new and alternative
explanation, but rather the old one that by the mid 1940s hehad previously ‘discarded’ (Myers 1951: 11, 1954b: 38).
For fishes, the old explanation had been best developed byCharles Tate Regan (1878–1943) at the British Museum(Natural History) in the year (1922) that Myers began hisvolunteer work at the American Museum of Natural History,his physical entry into the New York School. At this time hepresumably began to learn of ‘the strength mustered againstthem’ (Myers 1951: 11, 1954b: 38). This ‘strength’ heportrayed as a list of references: Matthew 1915; Noble 1925;Dunn 1931; Schmidt 1943; Simpson 1939, 1940a, 1940b;
Darlington 1948; Berry 1928; Chaney 1940. ‘Them’ meanspersons—such as Regan, destined in 1927 to become thesixth Director of the museum in South Kensington (Burneand Norman 1943)—who favoured, among otherexplanations for various organisms, the old explanation forfishes: ‘that in early Cretaceous times S. America and Africaformed one continent, which must have extended toIndia...The alternative view, that the Ostariophysi originatedin the north and spread southwards, involves so manyimprobabilities as to be almost unbelievable.’ (Regan 1922:206–207).
Using the phrase ‘strictly fresh-water’, Regan hadconsidered the same fishes (Ostariophysi) under the samepoint of view later adopted by Myers, who used the term‘primary’ rather than ‘strictly’. Regan’s (1922) paper waslater promoted as an ‘antidote to this influence’ [of the NewYork School] by one of Myers’ own students (Gosline 1944:213) in a paper that Myers (1949: 317) once cited as anexample 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 theassumed difference in the fishes’ physiological tolerance todissolved salt (also Gosline 1975).
A difference in physiological tolerance is fundamental towhat Myers considered his ‘one discovery worth generalquoting’ (see above). The discovery stemmed, seemingly,from experience in keeping aquarium fishes. It existed alsoas hope and expectation that were never realised: ‘I hope(and rather expect) that…physiological work…willdemonstrate a real basis for the primary and secondarydivisions’ (in litt. to Darlington, 14 July 1948). For him a realbasis would explain ‘why [primary fishes] are confined tofresh water’ (Myers 1949: 318) and justify the claim, ordiscovery, of their ‘ancient physiological inability to survivein salt sea water’ (see above). In his view this real basis, oncedemonstrated, would lie within physiological ecology, not ingeographic distribution itself, a fact neither obvious norwidely appreciated: ‘I am perfectly aware that most of thesecondary groups behave almost indistinguishably fromprimary ones in their fresh-water dispersal’ (in litt. toDarlington, 14 July 1948), yet, paradoxically, ‘it is…difference in [geographic] pattern…which has convinced methat the [six] groupings [primary, secondary, vicarious,complementary, diadromous, sporadic] have considerablebasis in fact.’ (Myers 1949: 320).
Madam How and Lady Why
The coherence and influence of the New York School are toopervasive, throughout much of the 20th century, to remainunexplained or dismissed (Menard 1986: 84) even as‘sophistry’ (Carey 1988: 133) or ‘solecisms’ (Carey 2000:129). One suggestion, to have been ‘wrong for the rightreasons’ (Laporte 1985, 2000), explains nothing about anymistake made in the interpretation of the biogeographic data,
Vicariance biogeography and the New York School 401
how ‘invalid’ inference derives from ‘valid’ principle(Laporte 1987: 331). Rather, the suggestion implies thatthere is no mistaken principle to understand and to explain—that to any question of earth history the biological data areeither neutral (Oreskes 1999: 295) or they are incompleteand, therefore, unreliable (Frankel 1981, 1984; cf. Craw1984c). If so, what then is the right reason, or valid principle,underlying Simpson’s (1966: 10) conclusion, based on datathen judged reliable and complete enough, that for example‘Australia has been in the fullest sense a dead-end’? There isno principle, none at all, beyond that of centre of origin.
If ideas about centres of origin are method-generatedartefacts of paralogous comparison, then the ideas and theirmistaken basis, even Simpson’s, are to that degree rationallyexplained – the how but not the why (Kingsley 1870). Yettheir artefactual nature was evident long before notions ofgeographic paralogy came into being (Cain 1944; Croizatet al. 1974; Nelson and Platnick 1984b).
The only other explanation suggested (by Croizat 1981:503) is that of Charles Darwin (1859: 352): ‘the simplicity ofthe view that each species was produced within a singleregion [centre of origin] captivates the mind’. So it did. Andso it does still, until one learns better and moves on.
Acknowledgments
For comment, information and literature we are grateful toGeorge Ball, Barbara Brown, Joel Cracraft, ChangMeemann, Daniel Cohen, Robin Craw, Jorge Crisci, MichaelCrisp, Andrew Drinnan, Malte Ebach, Gregory Edgecomb,Lance Grande, John Grehan, Michael Heads, WolfgangHennig, Marianne Horak, Christopher Humphries, PhilippeJanvier, David Johnson, Sylvia Kirchengast, Peter Linder,Jorge Llorente Bousquets, Malcolm McKenna, StephenMcLoughlin, Paul Manos, Bernard Michaux, Juan Morrone,Charles Myers, Lynne Parenti, Norman Platnick, Jay Savage,Scott Schaefer, Randall Schuh, Victor Springer, DennisStevenson, Jens Sommer-Knudsen, Pascal Tassy, FrançoisVuilleumier, Jonathan Waters and David Williams. We aregrateful also to William Cox and the Smithsonian InstitutionArchives for photocopies of correspondence betweenGeorge Myers and W. D. Matthew, P. J. Darlington andW. K. Gregory.
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