Palaeogeography, Palaeoclimatology, Palaeoecology
Permian Gondwana–Boreal antitropicality with special reference to brachiopod faunas
Introduction
Among all disjunct distributional patterns of modern biota, the north–south intercontinental disjunction, here referred to as antitropical distributions, is perhaps one of the first recognised but remains one of the least understood and most disputed biogeographic phenomena. In this context, antitropicality is defined as a pattern of biotic distribution in which forms of life ``shun the Tropics'' (Hubbs, 1952, p. 325). In this paper, we prefer the term antitropicality or antitropical distribution to an alternative term, bipolarity or bipolar distribution. The latter has also been used in the literature (e.g. Berg, 1933; Ekman, 1953; Ustritskiy, 1974), but as Hubbs correctly argued and also will be revealed in this paper, examples of truly bipolar distributions, namely life forms that are confined to the high-latitude regions of both hemispheres, are rare. Many previously cited references to the so-called bipolar distributions (s.l.) actually embrace a complex array of taxa whose geographical ranges may extend from the subtropical through temperate zones to polar regions in both hemispheres but are entirely absent from the intervening tropical belt. Other terms with a more or less similar meaning to antitropicality include antiequatorial distribution (Randall, 1981), bihemispherical (Whatley and Ballent, 1994), amphitropicality (Raven, 1963; note that this term literally means `two tropical', therefore should not be used in cases of true antitropical distributions, see Cox, 1990 for more discussion), and extra-tropicality (Astaf'yeva and Astaf'yeva-Urbaytis, 1992), but they all lack a precise definition and are therefore not followed here.
Antitropicality is very pronounced in the modern biosphere. It has been noted for almost all animal and plant groups, terrestrial and marine (see summary and reviews in Ekman, 1953; Darlington, 1957; Briggs, 1995). However, it is perhaps most prominent and most extensively documented in shallow marine faunas, especially near-shore fishes (e.g. Berg, 1933; Hubbs, 1952; Randall, 1981; White, 1986; White, 1994; Howes, 1991) and benthic molluscs (Lindberg, 1991; Crame, 1993 and references therein provided).
As a widely recognised biogeographic feature, antitropicality is of course not unique to the modern biosphere. This significant intercontinental disjunction has also been well documented in the geological record although its strength has varied from time to time judging by the number of taxa which demonstrate antitropical distribution (Fig. 1). Apart from a possible case of an Ordovician marine benthic antitropicality discussed by Fu (1998), few records of antitropical distributions are known prior to the Late Carboniferous. The Permian and Jurassic, on the other hand, appear to be the two geological intervals when antitropicality manifested itself most prominently, at a level that may be compared with or exceed that of the modern marine biosphere (Fig. 1).
While there is little doubt on the existence of antitropicality as a significant biogeographic pattern of the Recent biota, no consensus has been reached about the mechanism and timing of its formation. At least six different scenarios have been proposed to explain the origin of modern antitropicality (see Fig. 2). Crame (1993) has grouped these views into two broad, opposing scenarios: dispersal vs. vicariance. Historically, antitropical patterns and indeed all disjunct distributional patterns of biota have been interpreted as a consequence of natural dispersal (Darwin, 1859). In this model, antitropical disjunctions are thought to have resulted from long-distance dispersal of larvae or juveniles (in the case of shallow marine invertebrates) or spores and pollen from one hemisphere to another without leaving recognisable evidence of dispersal in the tropical region. However, how exactly larvae or spores/pollen are dispersed across the Tropics is still poorly understood. Several transtropical dispersal mechanisms have been suggested, including `island-stepping' (Fig. 2C), tropical submersion and coastal upwelling (Fig. 2D). Two key factors that need to be considered in adopting these `dispersalist' models are the availability of migration pathways at a given time and the migration potential of organisms, which in turn is a function of larval longevity, climatic regimes and (palaeo-)oceanographic conditions. The dispersalist school of views has appealed to many researchers and appears to remain one of the popular solutions to the antitropicality dilemma.
Almost parallel to the development of the dispersalist views, a different school of opinion argued strongly in favour of the vicariance approach (Nelson and Rosen, 1981; Briggs, 1987; Whiley, 1988). This hypothesis assumes that antitropicality may have originated as a result of an originally contiguous fauna or flora, later halved into distinct hemispheres by tropical extinction due to either excessive warming or competition, or by tectonic effects. The Relict Theory of Théel (1911) is perhaps the most well known and still the most widely accepted scenario within this school (Berg, 1933; Hubbs, 1952; Valentine, 1984). The Relict Theory basically interprets the antitropical disjunctions as relicts of a once cosmopolitan biota that existed during a globally more equable climatic regime (Fig. 2A). Using this hypothesis, the antitropical patterns of modern biota have been considered to have originated in accordance with the Pleistocene glaciation and global cooling event during which high-latitude taxa could migrate across the then cooling tropical region to form first a cosmopolitan pattern, but subsequent deglaciation and associated tropical warming extinguished those taxa that had migrated to the tropical region during the cooling regime, resulting in antitropical disjunctions. Similar to the relict theory, migration of taxa out of the tropical region followed by tropical extinction due to excessive competition has also been advocated as a possible explanation of the origin of modern antitropical disjunctions (Fig. 2B). The `centre of origin' hypothesis has been especially referred to the antitropical patterns in the Indo–western Pacific region, in that antitropical taxa are considered to have evolved from the warm tropical region and later migrated to the middle and high latitudinal regions due to excessive warming or competition (Briggs, 1987).
The third opinion on the origin of antitropicality ascribes the apparent morphological resemblance of so-called antitropical taxa to the effect of convergent evolution or convergence of morphological characters under the influence of similar ecological conditions (Fig. 2E). This argument, usually described as the parallelism view, has been especially favoured by botanists and palaeobotanists (Meyen, 1971; Asama, 1985; Oshurkova, 1996).
In this paper, we present data on the antitropical distributions of Permian Brachiopoda based on our independent as well as joint examinations of specimens from both the Arctic region (Arctic Canada, northern Russia) and Gondwanaland (Australia and Tibet) and Gondwanaland-derived terranes (e.g. Shan-Thai Terrane in SE Asia). The joint examination of the fossil material concerned was conducted when G.R. Shi visited Russia in 1995 and T.A. Grunt visited Australia in 1996. The fossils examined include both existing collections currently at the authors' disposal as well as material jointly collected by ourselves from the Pechora Basin of the northern Urals (in 1995) and the southern Sydney Basin of eastern Australia (in 1996). Although our focus is primarily on brachiopods, owing to their abundance and ubiquity in Permian shallow marine sedimentary rocks and our expertise on the taxonomy and biostratigraphy of this group, necessary references and discussions are also extended to other Permian fossil groups where appropriate data or literature is available.
For the first time, we have been able to examine and illustrate a range of Permian brachiopod taxa from both the Boreal and Gondwanan Realms in an attempt to assess the degree and nature of their taxonomic resemblance (including some long-acclaimed antitropical taxa). Earlier observations on the phenomenon of Permian antitropicality have been made by Fredericks (1931), Likharew (1960), Waterhouse, 1968a, Waterhouse, 1968b, Waterhouse, 1970, Waterhouse, 1971a, Waterhouse, 1971b, Ustritskiy (1974), Astaf'yeva (1991), Astaf'yeva and Astaf'yeva-Urbaytis (1992), Palmieri et al. (1994), Shi et al. (1995), Shi and Archbold (1995), and Shi and Zhan (1996), but few have explored the mechanisms responsible for the origin of the phenomenon. In this paper, in view of the data reviewed we argue for a plurality of possible mechanisms, migration pathways and timings for the Boreal–Gondwanan biotic interchanges during the Permian.
Section snippets
Definition and classification of Permian antitropicality
As the definition of antitropicality involves the distribution of certain taxa in space and time and with respect to patterns of climatic zonation, it is necessary to review, briefly, the Permian global marine biogeography and palaeogeography in which these antitropical patterns occurred (for recent, more detailed reviews on Permian global marine biogeography, see Grunt, 1995, Shi et al., 1995, Grunt and Shi, 1997, Jin and Shang, 1997, Shi, 1998). In general, three broad biogeographic realms
Comparison of stratigraphic ranges of antitropical genera
Although Permian antitropical species have been increasingly recognised, our observation of the Permian antitropicality remains largely based on genera. For this reason, here we only refer to the first and last occurrences of some selected genera of Brachiopoda for discussion (Fig. 5). In doing so, it should be born in mind that direct correlation between the Permian marine sequences of Gondwanaland and the Arctic region is hampered by strong Permian marine provincialism and is based on limited
Migration pathways and possible mechanisms
General scenarios with respect to the origin of modern antitropicality have been noted above. As far as the Permian antitropicality is concerned, several interpretations exist and may be grouped into two general schools for the convenience of discussion: migration (teleplanic migration or vicariance migration) vs. parallel evolution (or convergent evolution).
The view of parallelism, or homoplasy as otherwise known, has been particularly favoured by palaeobotanists presumably because they deal
Discussion and conclusions
A comparison of the Permian antitropicality with that of other, younger geological periods (e.g., Late Jurassic and the Recent) appears to suggest two common features: (1) that all had continents located at or near the poles; and (2) that a moderate- to high-latitudinal thermal gradient prevailed during these periods. It therefore seems suggestive to us that polarised continental positioning in both hemispheres and comparable cool to cold climatic conditions between the middle- and
Acknowledgements
This paper is the distilled version of oral presentations presented at several recent international conferences. On these conference occasions, we had opportunities to exchange information and learn from colleagues about the complex issues related to the Permian antitropicality. In particular, we would like to thank the following for discussions: N.W. Archbold, H.J. Campbell, J.M. Dickins, C.B. Foster, V.G. Ganelin, B.F. Glenister, Jin Yugan, S.S. Lazarev, T.B. Leonova, I.N. Manakov, Shuzhong
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