Research paperAnalytical methods in palaeobiogeography, and the role of early vertebrate studies
Introduction
Middle Palaeozoic fossils (e.g., trilobites, crustaceans) have featured in several recent publications concerning the use of fossils in biogeography (e.g., Lieberman and Eldredge, 1996, Ebach and Edgecombe, 2001, Rode and Lieberman, 2005). Largely overlooked, however, has been the key role of early vertebrate studies in the development of analytical methods involving cladistics and vicariance, and the application (use and misuse) of fossil data in biogeographic analysis (e.g., not even mentioned in the book by Lieberman, 2000). Siluro-Devonian vertebrates are morphologically complex, taxonomically diverse, and globally distributed fossils in both non-marine and marine aquatic environments, and known from numerous and a rapidly increasing number of localities (e.g., Janvier, 1996, fig. 8.6; Young, 2003, fig. 1). Some groups show empirical data with clear distributional differences between major taxa (Janvier, 1996, fig. 7.6), demonstrating their importance for biogeographic analysis.
This brief overview summarises the contribution of early vertebrate studies to research on palaeobiogeography, from the viewpoint that Siluro-Devonian vertebrates provide an unequalled dataset with which to assess the contribution of palaeontology to biogeography, and in turn the potential contribution of cladistic methodology to aspects of earth science, in particular the reconstruction of past geography (palaeogeography). In addition, I have tried to summarise the underlying logic in some seminal early papers concerning ‘centres of origin’ and dispersal hypotheses, topics that have been hotly debated in the recent literature as explanations of disjunct distributions in the modern biota. This debate is relevant to early vertebrate studies because many researchers now endorse the view that certain Siluro-Devonian vertebrate groups evidently originated in restricted areas, and later expanded their range by ‘dispersal’ into new areas, as evidenced by detailed analysis of their fossil records in time and space.
Just over 40 years ago, early vertebrate input was stimulated by Lars Brundin's presentation to the 4th Nobel Symposium on Hennig's methods of phylogenetic analysis, published in the volume ‘Current Problems of Lower Vertebrate Phylogeny’ (Ørvig, 1968). A range of early vertebrate papers in that volume included Gareth Nelson's analysis of acanthodian gill arch structure (Nelson, 1968). Inspired by the ideas of Hennig and Brundin, Nelson followed this with two short but highly significant papers providing insightful and far-reaching comments on historical biogeography in relation to phylogenetic analysis (Nelson, 1969, Nelson, 1970; for recent comment see Williams and Ebach, 2004). Phylogenetic methods, developed by entomologists (Hennig, 1950, Hennig, 1966, Brundin, 1966, Brundin, 1968) to deal with the most diverse living group (insects), had already been applied to the most diverse living vertebrates (teleost fishes) by Greenwood et al. (1966), so further elaboration by other ichthyologists was not unexpected.
Section snippets
Hennig's Progression Rule and Nelson's rule
Biogeographic applications were based on Hennig's (1966, pp. 133–139) ‘Progression Rule’, which employed a parsimony criterion (minimum dispersal) to estimate ancestral distribution and dispersal history by mapping geographic distributions onto a cladogram. Nelson (1969) discussed the method using an example of two sister species (C and D) occurring in two separate areas (x, y). Without additional evidence, the most parsimonious assessment of the distribution of their hypothetical common
Use of fossils in biogeography
Nelson (1969, p. 243) had considered the problem of historical biogeography to include the issue of incorporating evidence from both fossils and geology, but he noted that ‘regrettably, theories of past geological events often have had such appeal that biogeographers simply … interpreted their data accordingly’. A parsimonious hypothesis based on the distribution of extant species was theoretically testable using data concerning fossil representatives of the group, but a major problem concerned
Vicariance and cladistic biogeography
From the perspective that Hennig's Progression Rule involved an unnecessary assumption that dispersal and speciation must be associated (see Section 2), Platnick and Nelson (1978) developed the ideas underlying the alternative explanation of vicariance, into what can be termed ‘cladistic biogeography’. This provided a means of using distributional data for resolving a pattern of interconnections between areas, but without claiming to specify the nature of those connections. Platnick and
Converging area cladograms and ‘geo-dispersal’
Young (1981, p. 233) and other authors had already noted that ‘Vicariance Biogeography’, advocated as a preferred general explanation for the distribution patterns observed in the modern biota, was entirely dependent on the Mesozoic to Recent segment of geological history, which involved the breakup of Gondwana. In contrast, global palaeogeography for much of the Palaeozoic is interpreted as involving mainly continental collision, and the occlusion of intervening oceans. Under that
Cladistic analysis of geological/geophysical data
Rosen (1978) had suggested that the evidence of historical geology could be organised hierarchically, to facilitate the testing of any biogeographic pattern (whether caused by biotic dispersal, or by vicariance). However, not all such data can be treated in this way. Hierarchical structure may seem to be inherent in certain types of information (e.g., the ‘tree of life’, perhaps even the ‘universe in general and nature in particular’; Lieberman, 2000, p. 6). However, for the types of evidence
Conclusions and summary
Almost all previous discussions about the evidential basis from non-biological data sets (geology, geophysics) used to reconstruct past geography, and how such reconstructions are tested using biological data, have focussed on the respective values of historical compared to experimental, or quantitative compared to qualitative data. Completely overlooked has been the key question about the structure of the evidence being analysed (whether hierarchical or not), which then predicates the type of
Acknowledgments
This research has been a contribution to IGCP Project 491, and is supported by ARC Discovery Grant DP0558499 to G.C. Young and J.A. Long (‘Australia's exceptional Palaeozoic fossil fishes, and a Gondwana origin for land vertebrates’). Provision of facilities in the ANU Research School of Earth Sciences is acknowledged. For lengthy discussions on questions of biogeography and palaeogeography over many years I acknowledge numerous colleagues, including Alain Blieck, Daniel Goujet, Philippe
References (90)
Devonian palaeontological data and the Armorica problem
Palaeogeography, Palaeoecology, Palaeoclimatology
(1987)Early vertebrates and palaeogeographic models
Geobios
(1995)Application of cladistics to terrane history–parsimony analysis of qualitative geological data
Journal of Southeast Asian Earth Science
(1995)- et al.
Devonian macrovertebrate assemblages and biogeography of East Gondwana (Australasia, Antarctica)
Palaeoworld
(2010) - et al.
Une mise au point sur la géologie des continents issus du Gondwana
Annales de la Société Géologique du Nord (2nd s.)
(1994) Ancestral areas: a cladistic reinterpretation of the centers of origin concept
Systematic Biology
(1992)Ancestral areas: optimisation and probability
Systematic Biology
(1995)Parsimony analysis in historical biogeography and coevolution: methodological and theoretical update
Systematic Biology
(1990)Transantarctic relationships and their significance, as evidenced by chironomid midges, with a monograph of the subfamilies Podonominae and Aphroteniinae and the austral Heptagyiae
Kunglica Svenska Vetenskapsakademiens Handlingar
(1966)Application of phylogenetic principles in systematic and evolutionary theory
Croizat's panbiogeography versus phylogenetic biogeography
Hierarchical representation of hypotheses of homology
Geodiversitas
Devonian tetrapod from western Europe
Nature
Centres of origin and related concepts
Systematic Zoology
The resurrection of oceanic dispersal in historical biogeography
Trends in Ecology & Evolution
A new wuttagoonaspid (Placodermi, Arthrodira) from the Lower Devonian of Yunnan (South China): origin, dispersal, and paleobiogeographic significance
Journal of Vertebrate Paleontology
The earliest phyllolepid (Placodermi, Arthrodira) from the late Lochkovian (Early Devonian) of Yunnan (South China)
Geological Magazine
Cladistic biogeography: component-based methods and paleontological application
The use of paleontology in systematics and biogeography, and a time control refinement for historical biogeography
Paleobiology
Vicariance biogeography
Phyletic studies of teleostean fishes, with a provisional classification of living forms
Bulletin of the American Museum of Natural History
Areas of endemism: definition and recognition criteria
Systematic Biology
Grundzuge einer Theorie der phylogenetischen Systematik
Phylogenetic Systematics
Abundance and diversity of labyrinthodonts as a function of paleolatitude
American Journal of Science
Early Vertebrates
A new Groenlandaspidid Arthrodire (Vertebrata: Placodermi) from the Famennian of Belgium
Geologica Belgica
Hyrcanaspis bliecki n. g. n. sp., a new primitive euantiarch (Antiarcha, Placodermi) from the Eifelian of northeastern Iran, with a discussion on antiarch phylogeny
Neues Jahrbuch fur Geologie und Palaontologie Abhandlungen
Devonian rhizodontids and tristichopterids (Sarcopterygii: Tetrapodomorpha) from East Gondwana
Transactions of the Royal Society of Edinburgh: Earth Sciences
Early Cambrian paleogeography and tectonic history: a biogeographic approach
Geology
Testing the Darwinian legacy of the Cambrian radiation using trilobite phylogeny and biogeography
Journal of Paleontology
Paleobiogeography: Using Fossils to Study Global Change, Plate Tectonics, and Evolution
Paleobiogeography: the relevance of fossils to biogeography
Annual Review of Ecology and Systematics
Trilobite biogeography in the Middle Devonian: geological processes and analytical methods
Paleobiology
Systematics and biogeography of the ‘Malvinella group’, Calmoniidae (Trilobita, Devonian)
Journal of Paleontology
A possible ‘elpistostegalid’ fish from the Devonian of Gondwana
Proceedings of the Royal Society of Victoria
A new buchanosteid arthrodire (placoderm fish) from the Early Devonian of the Ural Mountains
Journal of Vertebrate Paleontology
On the identification of areas of endemism
Systematic Biology
Historical biogeography: introduction to methods
Annual Review of Ecology and Systematics
Gill-arch structure in Acanthodes
The problem of historical biogeography
Systematic Zoology
Outline of a theory of comparative biology
Systematic Zoology
Comments on Leon Croizat's biogeography
Systematic Zoology
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