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The Early Devonian palaeobiogeography of Eastern Australasia
Palaeogeography, Palaeoclimatology, Palaeoecology 444 (2016) 39–47
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The Early Devonian palaeobiogeography of Eastern Australasia Elizabeth M. Dowding ⁎, Malte C. Ebach School of Biological, Earth & Environmental Sciences, UNSW, Kensington, NSW 2052, Australia
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Article history: Received 27 August 2015 Received in revised form 6 November 2015 Accepted 24 November 2015 Available online 12 December 2015 Keywords: Australia Biochorema New Zealand Parsimony analysis of endemicity Tectonostratigraphic terranes
a b s t r a c t Biogeographical analyses of Devonian Australasia (Australia and New Zealand) have previously presented a narrative approach to the interrelationships of continental faunal provinces. Useful for the creation of biotic hypotheses, similarities and differences between biotic provinces remains untested. This study seeks to propose the first regional faunal provinces allowing a test of geological and biotic hypotheses using Parsimony Analysis of Endemicity. The relationships of eastern Australasian terranes were tested through the hierarchical analysis of faunal composition. Biochorologic sub-provinces were proposed using geo-spatial computer software and analysed through phylogenetic software that resulted in the formalisation of 20 biochorologic sub-provinces. The relationships between these units show a significant holdover of Silurian species, the interactions between convergent terranes and the continental margin, the distinct history of the Lachlan Fold Belt terranes, and offers biotic support for the existence of the Tasmanian microcontinent Vandieland. © 2015 Elsevier B.V. All rights reserved.
1. Introduction Encompassing 70% of the continental area, Devonian Gondwana saw one of the earliest major adaptive radiations in Earth's history, giving rise to many evolutionary important taxa (Scotese et al., 1999; Talent et al., 2000; Boucot et al., 2013). With a collection of tectonostratigraphic terranes (‘terranes’, fragmentary crustal segments sutured to a continental margin presenting unique and distinctive geological history), subduction zones, and back-arc and forearc basins, Gondwana's Early Devonian presented a dynamic environment and myriad habitats (Fig. 1, McElhinny et al., 2003). Proposed by Young (1981), Eastern Gondwana, encompassing Australasia, forms the Wuttagoonaspid–Phyllolepid faunal province. This province stands distinct from the remaining four provinces: the Cephalaspid province (Euramerica), Amphiaspid province (Siberia), Tannuaspid province (Tuva), and Galeaspid–Yunnanolepid (South China) (Young, 1981). Based primarily upon fish fauna, biogeographical studies describing continental relationships have also been made using brachiopods (Talent et al., 1972; Yolkin et al., 2000), conodonts (Talent et al., 1972), radiolarians (Afanasieva and Amon, 2013), gastropods (Cook, 2001), trilobites, sponges and polyzoans (Talent et al., 1972) respectively. Single taxa studies, however, inevitably leave informational gaps which prove problematic considering that associations and affinities drawn are clouded by the individual group's evolutionary history and relationships. Furthermore, for the creation of true representations of biotic areas, data from multiple taxa types, habits, and life histories ought to be incorporated; the patterns and conclusions drawn from ⁎ Corresponding author. E-mail address: [email protected] (E.M. Dowding).
http://dx.doi.org/10.1016/j.palaeo.2015.11.037 0031-0182/© 2015 Elsevier B.V. All rights reserved.
single taxa studies inherently missing significant palaeobiogeographical patterns (Damborenea and Macenido, 1992). A solution is to incorporate the distribution of multiple unrelated taxa in order to determine biotic overlap. Biochorologic units, such as biochorema and biochorologic sub-provinces, are endemic biotic centres within a geographic envelope alter in time, distribution, and scale, offering spatial and temporal analysis of endemism and distribution (Cecca and Westermann, 2003). Such an approach is seen to give the best coverage of the habited area, relieving in part absence signification due to the specimen's inability to survive geological action or fossilisation (Aldridge and Purnell, 1996; Girard and Renaud, 2011). For example, the coastline of the Early Devonian Australian margin is based upon the zonation of the conodonts Pedavis pesavis and Icriodus woschmidti (Mawson and Talent, 2003; Fig. 2). The ubiquitous nature of conodont deposits have seen them used as identifiers for biostratigraphy (Lyons and Percival, 2002; Mawson and Talent, 2003; Girard and Renaud, 2011). However, within the convoluted frame of a convergent back-arc basin, the use of a single taxon maintains its difficulties. Marine transgressions were a known feature of the Australian margin, supplemented by continuous marine deposition within a back-arc and the Wagga Marginal Sea, an ambiguity left unresolved by Talent et al. (2000) (Glen and Walshe, 1999; Glen, 2013). Whilst their palaeoecology makes no part of biochorologic characterisation, the formation of faunal sub-provinces will logically display the patterns of their deposition. The proposal of Talent et al. (2000) is contradicted by the presence of the marine genus of brachiopod Chonetes within deposits from the Wagga–Omeo terrane (VandenBerg et al., 1976). Endemic centres created through data gained from multiple taxa are an established approach for the hierarchical analysis of areas (Cecca and Westermann, 2003). These centres are bound by the
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Fig. 1. Devonian Gondwana, area of study highlighted. Reconstruction from Boger (2011).
temporary range limits of the constituent taxa, ranks scaling with the degree of endemism, taxa distribution, and duration (Westermann, 2000). The Early Devonian Australasian coast, presents a myriad of environments respectively affected and disrupted by terrane convergence and the shifting of the Gondwanan supercontinent (VandenBerg et al., 2000). During the Early Devonian, Gondwana was steadily moving towards Panthalassa, its active margin interacting with a geological collage of terranes and fragmentary blocks (Scotese et al., 1999; von Raumer and Stampfli, 2008; Fergusson, 2010; Boucot et al., 2013). Obduction, followed by subduction inversion formed the Early Devonian palaeotethys between the Gondwanan and Variscan terranes, East Gondwana proving convergent in a back-arc setting (Von Raumer et al., 2002; Fergusson, 2010; Boger, 2011). This movement, coupled with an alteration in the locus of convergence within Gondwana, set in motion a complex history of accretion, rifting, strike-slip faulting (Siluro-Devonian), and extensional/contractional phases across its eastern portion (Foster and Gray, 2000). During the course of this convergence, the geography and fauna are seen to have changed and altered with the geology (Lieberman, 2005). Silurian holdover faunas were preserved into the Early Devonian by the Wagga Marginal Sea and the rift and extensional basins that riddled the Australian coast (Talent et al., 1972). These Silurian fauna transitioned into more typical Devonian community compositions contemporaneously with the westward accretionary movements of the outboard terranes
(Veevers, 2004; YuanSheng et al., 2008). The nature of the relationships between these two coeval changes can be hierarchically represented, regional faunal provinces or ‘sub-provinces’ identified and their geological relationships inferred. Contrary to biochorologic and terrane models, tectonic models for Australasia are well established; the timing and nature of terrane accretions and basin formations being generally resolved within the Early Devonian (Cawood, 2005; Fergusson, 2010; Boger, 2011; Fergusson and Henderson, 2015). The Tasmanides are characterised by the Lachlan Fold Belt and Thomson Orogen, age of accretion younging from west to east (Foster and Gray, 2000; Glen et al., 2004). The central Lachlan Fold Belt, comprised of Howqua–Tabberabbera and Wagga-Omeo, exhibits an interesting correlation to the study of Yolkin et al. (2000). Analysing brachiopod community compositions, Yolkin et al. (2000) aligned the central Howqua–Tabberabbera and Molong–Monaro terranes of the eastern Lachlan Fold belt as a single biotic province depicting relation to Melbourne and New Zealand (Fig. 3). The constituent terranes of New Zealand were not delineated by Yolkin et al. (2000) and the relationships of these, consequently, will be sought here though the continuity of multiple taxa and bound by established terranes. The Melbourne, Tasmanian, and New Zealand terranes contain some of the most richly fossiliferous depositions within Australasia. Recently they have come under much scrutiny, their accretionary history generating many hypotheses such as ‘Vandieland’ (Cayley et al., 2002;
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Cayley, 2011), the reconstruction of Li and Powell (2001), and the placement of New Zealand between Australia and Antarctica (Ireland, 1992; Fagerstrom and Bradshaw, 2002). Described as a continental fragment, magmatic analysis of eastern Gondwanan granitoids have indicated that those of New Zealand, Northern Victoria Land, Marie Byrd Land in West Antarctica, Tasmania, and central Lachlan Fold belt (Howqua–Tabberabbera and Wagga-Omeo) were correlated, potentially representing a semicontinuous magmatic belt in excess of 2000 km along the Gondwanan margin, prior to subsequent rifting and breakup (Cooper and Millener, 1993; Daugherty et al., 1993; Muir et al., 1996). Evidence found within New Zealand's Reefton group, (the distribution and evolutionary history of endemic species), give biotic supplement to the established links between Tasmanian, Victorian, and Antarctic sedimentary basins (Fagerstrom and Bradshaw, 2002; Adams et al., 2013). The aim of this study is to propose the first Early Devonian biochorologic sub-provinces, presenting their hierarchical relationships to discern the affinities and geological relations of tectonostratigraphic terranes within Australasia. Focussing upon the Australasian coastline during the early Devonian (419–393mMa), this study will apply Parsimony Analysis of Endemicity (PAE) to existing hypotheses of terrane history (Young et al., 2010), palaeogeography (Talent et al., 2000) and biotic area evolution (Ebach and Edgecombe, 2001). Using faunal and biochorologic data, new regional configurations of Australasian geological and biotic associations will be proposed. This paper will also provide supplementary evidence for the existence of Vandieland, the location of New Zealand, and establish the nature of the relationship between the terranes of the Lachlan Fold Belt. Devonian biogeographical reconstructions have largely focussed upon information from single taxa (e.g. genera and families). The patterns and conclusions are drawn from these analyses focusing upon the life histories of single taxa and inherently miss significant palaeobiogeographical patterns (Parenti and Ebach, 2009). Similarly, geographical or geological approaches to reconstruction remain limited in their ability to suggest faunal distribution and seaboard. This study seeks to understand the dualism between the evolution of location and life. 2. Materials and Methods 2.1. Biogeographic Setting Fig. 2. The Early Devonian marine seaboard of Australia proposed by Talent et al. (2000).
Early Devonian (419–393 Ma) Australasia encompassed Australia and New Zealand respectively, New Guinea not yet having been formed.
Fig. 3. Faunal relationships based upon brachiopods of Early Devonian crustal blocks (Yolkin et al., 2000).
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Located along the Gondwanan eastern coastline, Australasia exhibits 14 early Devonian tectonostratigraphic terranes and basins, the former being fault bound regions with a unique stratigraphic history (Leitch and Scheibner, 1987). These units are generally allochthonous and present dynamic histories of rift and suture, interacting with a convergent plate boundary (Cawood and Leitch, 1985). The 14 Devonian Australasian terranes and basins are used for analysis of geological evolution based upon their faunal characteristics. The relationships between these terranes will be assessed through the compositions of their biota. Selected terranes are chosen based on endemic taxa, in particular trilobites, brachiopods, fish, sarcopterygians, and placoderm faunas. The terranes were required to show some overlapping faunal distributions, and have been verging upon or accreted to the Gondwanan margin after the Early Ordovician and before the Early Devonian. 2.2. Data Collection The following data was collected manually through an extensive review of the existing literature, such as, the terrane maps of Leitch and Scheibner (1987) and Vandenberg et al. (2000) for Australia and New Zealand (see Appendix A, Table 1). The data includes all recorded brachiopods, trilobites, sarcopterygians, and placoderms from 14 terranes and basins. 91 Early Devonian (419–393 Ma) formations across 14 terranes and basins were described in terms of type, composition, and environment of deposition. The tectonostratigraphic terranes, being fragmentary, unique, lithological units, were required to have both endemic and more cosmopolitan inhabitant biota (VandenBerg et al., 2000). Devonian corals and conodonts were discounted due to their ubiquitous distributions in favour of brachiopods, trilobites, sarcopterygians, and placoderms. Patterns of endemicity and distribution would be clouded by the informational bias presented by the cosmopolitan nature of these taxa (Young, 1987). Accordingly, sarcopterygian, placoderm, trilobite, and brachiopod presences were recorded within formation and corollary terrane, data pulled from previous palaeontological works (Table 1). Genus, rather than species, was retained in order to account for disarticulation and the inability to specifically identify all specimens within fossil record, allowing a display of higher order relationships and patterns (Upchurch et al., 2002). The location of the specimen was noted in terms of latitude and longitude, and when the specific location was unavailable, the formations location was recorded. Taxon age was not incorporated to analysis; distribution, composition, and location used for character individuation (Young, 1987, 2007). 2.3. Terranes and basins The following terranes and basins were used in the PAE analysis: Buller Terrane. — Part of Devonian New Zealand the Buller Terrane comprises 11 fossiliferous formations. The formations of this terrane exhibit a high energy, near shore beach to shelf depositional environment. Canning Basin. — Used as an outgroup to root the tree. No formations were noted for the intercratonic basin, with low energy, deep-water to hypabyssal deposition. Darling Basin. — This basin was muddy, slowly subsiding, and asymmetrical; a shallow marine depositional environment with deltaic, fluvial sediments. Georgina Basin. — Cratonic Australian segment presenting two fossiliferous Devonian formations. Non-marine, low energy basinal palaeoenvironment. Hodgkinson–Camel Creek Terrane. — Fossils were found in a medium to low energy ocean floor and trench accretionary prism.
Howqua–Tabberabera Terrane. — This composite terrane is comprised of 4 individual formations and the Wentworth group. Howqua–Tabberabera exhibited a muddy bottomed, lacustrine to shallow marine depositional environment. Bioherms have been noted within the terrane, suggesting medium to low energy environments. Melbourne Terrane. — 17 formations are found to represent deep water palaeoenvironment. Medium to low energy environment can be seen through the transport channels and rippling that marks the terranes beds, laminations, and formations. Located at the back of the Kanmantoo orogeny, it was a foreland basin moving into orogenic deformation during the Middle Devonian Tabberabberan orogeny. Molong–Monaro Terrane. — This composite terrane exhibits traits of a shallow marine shelf. The 27 formations vary between low energy muddy substrates, volcaniclastics, and patch reefs complete with reef mounds and some megaturbidites forming within the forearc basin. Molong–Monaro presents basins and rifting, both of which were associated with terrane dispersal. The tectonic highs, between the troughs and basins, were progressively closed during the Devonian by the twin actions of the Tabberabberan and Bowning orogenic events. Narooma Terrane. — The Devonian segment of this allocthonous terrane is limited to the Bunga Beds. Situated against an escarpment within a volcanic graben, the Bunga Beds present deep water deposition, a freshwater lake dammed by the graben. The Narooma Terrane was accreted during the Benambran orogeny, presenting outer arc slope and trench during the early Devonian (Stokes et al., 2015). Sydney–Bowen Basin. — Volcanically active, the Sydney–Bowen Basin presents shallow marine and terrestrial sedimentation, succeeded by deeper water. High energy conditions gradually decrease with the increasing water depths into a foredeep. Tamworth–Yarrol Terrane. — This composite terrane (Wisemans Arms, Woolomin, Cockburn, and Texas terranes) encompasses a forearc basin in the Graveyard Creek foredeep and the Kangaroo Hills deep, with two superimposed arc fringe-forearc basin sequences. Extensively bioturbated, this terrane had a low energy environment characterised by siltstone and sandstone and some volcaniclastics with shallow water environments in the west found within six formations. North-West Tasmanian Terrane. — Though a low to medium energy environment, there is evidence of strong current activity within this terrane. Rapid deposition without much washing can be seen in this uplifting depositional interface which moved into shallow water. This pattern can be seen graded within each of the seven North-West Tasmanian formations. Takaka Terrane. — Steep eastward dipping has been noted within the Takaka terrane. Characterised by the Devonian Baton formation, Takaka is a mud rich, deep offshore environment with present turbidites. Wagga-Omeo Terrane. — This relatively low energy terrane presents a shallow marine depositional environment as seen across its 6 Devonian formations. During the Devonian the Wagga-Omeo Terrane underwent dispersal of the order of kilometres to tens of kilometres, rifting throughout the Early Devonian and Late Silurian.
2.4. Biochorologic sub-provinces Biochorema are endemic biotic centres within a geographic envelope and therefore alter in time, distribution, and scale (Cecca and Westermann, 2003). For this regional study, biochorema are referred
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to as sub-provinces. 20 biochorologic sub-provinces are proposed, characterised by 10% endemicity of fossil genera; the recommended unit for provincial studies (Guangrong, 1998; Anstey et al., 2003; Andreu et al., 2007). These biochorologic sub-provinces were identified using Biodiverse 0.19, a spatial analysis tool for the analysis of diversity, which distinctively offers the ability to map and analyse georeferenced biological specimens (Laffan et al., 2010). Within this analysis, clustered taxonomic dissimilarity relationships were used and areas and taxa that did not achieve 10% endemicity were removed. In this way, Biodiverse allows a distinctive insight to the similarity and dissimilarity of spatial diversity, displaying endemic centres and presenting relationships within a hierarchical phenogram. The recognition of a hierarchy within the results leads to the implication, following from Anstey et al. (2003), that smaller units may be nested within larger ones. The majority of biogeographical studies (Young, 1981; Rich and Young, 1996; Young et al., 2010) have presented large geographic areas, this study seeking to unbind the nested relationships seen within continental studies. This recognition of biogeographical hierarchy allows the discernment of how endemic regions or sub-provinces have been formed through ascertainable evolutionary process (Rosen, 1988; Rosen and Smith, 1988; Anstey et al., 2003). Because of the proximity of each of the deposits, differences seen between the sub-provinces are assumed to be related more to geological action and consequent habitat type as opposed to climatic influence (Yanin, 2010). To this end, polytypic areas prove especially advantageous, giving equal valency to all habitat types and capturing signals given by mixed or transitional zones and environments (Guangrong, 1998).
2.5. Analysis of tectonostratigraphic terranes Tectonostratigraphic terranes, being fragmentary crustal segments sutured to a continental margin presenting unique and distinctive geological history, were analysed by their biotic similarities. Proposed by Rosen (1988) and Rosen and Smith (1988) endemicity data, hierarchically structured within a parsimonious phenogram, can be used as a pluralistic approach to discerning geological and stratigraphic relations. The use of biotic data for area classifications has been widely utilised following Rosen's (1988) Parsimony Analysis of Endemicity (PAE). PAE defines endemic areas, using these to determine hypothetical relationships in the absence of phylogenetic data (Morrone and Escalante, 2002; Nihei, 2006; Morrone, 2014a). Based upon taxa presence-absence data matrices, PAE constructs parsimonious cladograms allowing the identification of ‘biotic components’: historical biotas, biogeographical assemblages, taxonomic assemblages, species assemblages, and their hypothetical relationships (Garcia-Barros et al., 2002; Morrone, 2014b). For each terrane, the genera profile was used to provide character individuation; distributional data entered as a presence ‘1’ only matrix and analysed using the parenthetical cladistics program LisBeth v.1.3 (Zaragueta Bagils et al., 2012). Noted within critiques, PAE groups on absences, thereby creating similarities between areas that share no taxa. Many have argued that this fails to capture historical complexity (Morrone and Escalante, 2002; Garzon-Orduna et al., 2008). Binary forms, such as PAE, often become problematic in the face of an incomplete fossil record, as relationships based upon sparse or well distributed data are at risk of losing presence significance to overwhelming sequenced absence data (absence signification). The use of LisBeth v. 1.3 within the present study, applies a parenthetical matrix, thereby allowing only presences. LisBeth v.1y3 was run using three item analysis and compatibility (3ia function), finding the most optimal (i.e. compatible) tree from the largest sum of fractional weighting (Nelson and Ladiges, 1996). Characters are depicted as hierarchies or polarized and ordered tree states, congruent or optimal tress summarised within the intersection tree (Zaragueta Bagils et al., 2012).
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Following from Young (1987), temporal information was seen to be an incorporated feature of the taxon or terrane, considered at terminal braches rather than nodes. This option is available due to the omission of phylogenetic data (Upchurch and Hunn, 2002), namely that distributions seen as indicative of geology rather than the specific evolutionary history of a taxon. 3. Results The apical tectonostratigraphic terranes (Fig. 4) correlate to the allochthonous VandieLand of Cayley (2011). The New Zealand terranes, Takaka and Buller, are proposed to have interacted prior to the accretion of Vandieland during the late Cambrian–Ordovician (Cooper and Millener, 1993). The central segment of the terrane tree intimates detail of the interactions of the Macquarie Arc and Narooma Accretionary Complex, assumedly falling central due to their shared history with the Howqua–Tabberabbera and Wagga-Omeo terranes (Table 1). The terrestrial, freshwater, mixed profile basins and terranes of Hodgkinson– Camel Creek and the Georgina Basin, basally group. Through the Biodiverse analysis of spatial cluster and endemicity, 20 biochorologic sub-provinces are proposed across the region, distribution allowing the proposed configuration of the Australian seaboard (Fig. 5), Molong-Monaro and Melbourne terranes offer the highest number of sub-provinces across time (Table 2). The analysis of the biochorologic subregions (Fig. 6) is seen to mirror those of the terranes (Fig. 4). The apex of the tree (Fig. 6) broadly reflects the terrane relationships of Fig. 4, grouping the endemic centres that fell within the Melbourne, Molong–Monaro, Buller, North-West Tasmanian, Howqua– Tabberabbera terranes, and the Sydney–Bowen Basin. Interestingly, the base of the tree indicates a temporal split, Pragian and Emsian faunal distributions grouping together, Lochkovian sub-provinces dominating the upper half of the tree. 4. Discussion Spatially and temporally bracketed 20 new biochorologic subprovinces give crucial insight to the biotic development of Eastern Australasia, its palaeogeography, and the dependence upon geological action through time (Fig. 5, Appendix A, Table 2). The interrelationships of these biochorologic units to their facilitating geology offer supplement to
Fig. 4. Analysis of terranes. The analysis of the terrane data returned one tree with a completeness (CL) of 92.0% and a retention index (RI) of 0.76.
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Fig. 5. a. Seaboard for the Early Devonian of Australia following the marine line of the Sydney–Bowen Basin and the biochorologic subprovinces. 1 — Mulga (Darling Basin), 2 — Dulcie (Georgina Basin), 3 — Cravens-Toomba (Georgina Basin), 4 — Ukalunda (Sydney–Bowen Basin), 5 — Humevale (Melbourne), 6 — Amphitheatre (Darling Basin), 7 — Boola (Melbourne), 8 — Florence (North-West Tasmania), 9 — Trundle (Wagga-Omeo), 10 — Walhalla-Kilgower (Melbourne and Howqua–Tabberabbera), 11 — Derriwong (Wagga-Omeo), 12 — Mandagery (Molong–Monaro), 13 — Cavan (Molong–Monaro), 14 — Garra (Molong–Monaro), 15 — Crudine (Molong–Monaro), 16 — Bowning (Molong–Monaro), 17 — Boyd (Narooma), 18 — Wurutwun (Melbourne), b. New Zealand.19 — Reefton (Buller), 20 — Baton (Takaka).
geologic models, providing evidence of the rotation of the Victorian region and Melbourne zone, the closure of the Wagga Marginal Sea, the nature of the relationship of continental basins and allochthonous terranes, and the biotic affinities of the three segments of the Lachlan Fold Belt. The analogous nature of the relationship between geologically based models and the results seen here confirm the efficacy of our palaeobiogeographic approach to regional biotic and geologic hypotheses. 4.1. Evidence for interaction between the Darling Basin the Lachlan and Thomson Orogens The Darling Basin covers two Biochorologic sub-provinces, being the Emsian Mulga (Table 2, number 1) and the Lochkovian to Pragian Amphitheatre (Table 2, number 6). Previously given only perfunctory attention (Fergusson, 2010; Glen et al., 2013), the results have indicated
the Darling Basin was a dynamic region, influenced by the movements of the Lachlan and Thomson Orogens making it both a cosmopolitan hub (Amphitheatre sub-province) and subsequently a refuge basin (Mulga sub-province) (Fig. 6). The analysis indicates that the Amphitheatre sub-province had biotic exchange with the sub-provinces of the Wagga-Omeo terrane. During the Lochkovian, the Amphitheatre sub-province was flanking the Wagga Volcanic Arch, overlap sequences of the Cobar-supergroup deposited within the rifts and flanking shelves of both the Lachlan and Thompson Orogens (Webby, 1972; Glen et al., 1994; Fergusson, 2010; Glen, 2013). Initially afforded by a period of major extension within the region, the relationship between the Darling Basin and the Orogens is limited to the Lochkovian. Due to interplay between the orogens, the Cobar basin was obliquely opened and then closed forcing the cessation of the eastward flowing
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problematic reconstructions (Fig. 2). During the early Devonian the Wagga Marginal Sea was closed by the continuous rollback of the subduction hinge coupled with regional uplift and strike-slip faulting (Lambeck and Stephenson, 1986; Fergusson, 2010). These movements lead to increasing endemism amongst the LFB subprovinces, characterised by refuge basins (Crook et al., 1973; Crook, 1980; Strusz, 2000), and intermediate and freshwater environments. The taxa profile have drawn the seaboard for the Pragian (Fig. 5), the Wagga marginal sea shown to not be fully closed, limited by the mixed profile of the Cavan sub-province (Table 2, number 13). The depiction of the LFB as a single geological unit was to an extent mirrored in Yolkin et al. (2000), who used brachiopod genera to group the Lachlan Fold Belt terranes of Howqua–Tabberabbera and Molong– Monaro into a single biotic unit. This study, however, has shown that Howqua–Tabberabbera and Molong–Monaro not only have distinct biochorologic sub-provinces (Table 2), but also that these subprovinces are more closely related to those of the Melbourne terrane than either are to each other (Fig. 6). Furthermore, during the early Devonian the Wagga margin was subducting between Gondwana and the Molong Island arc, maintaining the separation of the Howqua– Tabberabbera and Molong Monaro terranes (Fergusson, 2010). Fig. 6. The analysis of biochorlogic sub-provinces. The tree presents a completeness of 75.3% and a retention index (RI) of 0.808.
palaeocurrent (Glen and Walshe, 1999). The termination of this current, coupled with the closure of the Wagga Marginal Sea lead to the isolation of Darling Basin, the fauna of the Amphitheatre sub-province transitioning into those of the Mulga. The Mulga sub-province is proposed here to have been a refuge basin, the persistence of late Silurian and Lochkovian fauna into to the Emsian apparent through the close relationship of the Darling Basin sub-provinces to each other than to another area (Table 2, Fig. 6). 4.2. The shared biochorologic sub-province of Hodgkinson–Camel Creek and Georgina Basin The Hodgkinson–Camel Creek terrane is one of Australia's youngest accretionary complexes (Withnall and Henderson, 2012). Formed within a Siluro-Devonian forearc accretionary prism the Cravens-Tooma sub-province (Table 2, number 3) exhibits the characteristics of a convergent margin with widespread, deep marine turbidites. The subprovince is, however, found to have been equally influenced by the continental margin (De Keyser and Lucas, 1968; Arnold and Henderson, 1976; Day et al., 1978). Constructed jointly out of the convergent margin and the continental freshwater of the Georgina basin, CravensToomba represents a composite sub-province (Table 2). Terrigenous materials and freshwater species were deposited into the marine Hodgkinson basin by strong, heavily laden currents from the west which lost most of their load and force as it moved towards the more distal parts of the basin (de Keyser and Lucas, 1968; Haines et al., 2001; Li and Powell, 2001). Whilst the exact linkages between the inland Georgina Basin and the convergent Hodgkinson–Camel Creek terrane cannot be directly ascertained, this sub-province does provide crucial insight into the interactions between inboard terranes and the Gondwanan margin. 4.3. The discrete history of the Lachlan Fold Belt terranes Often depicted as a single accretionary unit (Li and Powell, 2001; Boger, 2011), the terranes of the Lachlan Fold Belt exhibits a suite of unique biotic sub-provinces (Tab 2). These relationships between the sub-provinces of the Lachlan Fold Belt are often contained within the borders of their host terrane, biotic similarities offering insight into the southern extension of the LFB, a zone which has been prone to
4.4. Vandieland The Melbourne terrane exhibits four sub-provinces including Humevale, Boola, Walhalla-Kilgower, and Wurutwun (Table 2 number 7, 10, and 18). During the early Devonian the Melbourne zone formed a depositional trough, sediments from the Stawell, and Bendigo zones, Selwyn Block inclusive, providing sedimentary material (Fergusson, 2003; Cayley, 2011). This external influence upon the terranes subprovinces coupled within the strong affiliations between them and those of North West Tasmanian provide biotic evidence for Vandieland (Cayley et al., 2002; Cayley, 2011). Vandieland, composed of the Selwyn block, the Melbourne Zone, and Tasmania was proposed by Cayley et al. (2002) and Cayley (2011) to explain the trapped pre-Cambrian continental extension of Tasmania, being the Selwyn block, and its occurrence upon mainland Australia (Crawford et al., 2007; Berry et al., 2009; Moore et al., 2015). Rotation of the Melbourne zone, evidenced by the migrating axis of trough deposition, which moved eastward providing links to the late early Devonian Molong-Monaro sub-province Cavan, and influencing the closure of Wagga Marginal Sea (Webby, 1972; Lambeck and Stephenson, 1986; Miller and Gray, 1996; Cayley et al., 2002). The delineation of this microcontinent and its allochthonous history is understood to provide a unifying framework for the explanation of the ‘Selwyn paradox’, and the observed geological connections between Tasmania, New Victoria Land, and Victoria (Veevers, 2004; Cayley, 2011; Moore et al., 2015). 4.5. New Zealand Strong association between Baton, Reefton, Humevale, and Florence suggest a proximal relationship between NZ and the southern Australian margin (Fig. 6). Previous studies have indicated that New Zealand was located between the 30th and 40th palaeolatitidue (Muir et al., 1996; Fagerstrom and Bradshaw, 2002; Torsvick and Cocks, 2013), a location which would allow exchange between Australian populations and those previously observed within Antarctic sedimentary basins (Adams et al., 2013). Magmatic analysis of granitoids have indicated that those of New Zealand, Northern Victoria Land, Marie Byrd Land in West Antarctica, Tasmania, and the central Lachlan Fold belt (Howqua -Tabberabbera and Wagga-Omeo) were correlated, potentially representing a semicontinuous magmatic belt in excess of 2000 km along the Gondwanan margin (Cooper, 1989; Cooper and Tulloch, 1992; Daugherty et al., 1993; Muir et al., 1996). Tectonically activated variation, with particular regard to the extensional and contractional
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phases of the LFB, altered the nature of both sedimentation and habitat, consequently shifting affiliations of the Lachlan Fold Belt away from the uninterruptedly stable sub-provinces of Tasmania and New Zealand (Fig. 6). The placement of New Zealand between the 30th and 40th latitude is therefore seen to complement the temporal changes marked between these New Zealand sub-provinces and the Victorian terranes. Both the New Zealand and Tasmanian terranes were primarily unaffected by the contractional, extensional phases of the Lachlan Fold Belt and consequently their links to the aforementioned remain strongest within the earliest Devonian (Fig. 6). 5. Conclusion Devonian Gondwana saw one of the earliest major adaptive radiations in Earth's history, giving rise to many evolutionarily important taxa (Scotese et al., 1999; Talent et al., 2000). This study presents the first regional palaeobiogeographic analysis of Early Devonian Australasia across a variety of taxonomic groups. Through the use of these multiple taxa, 20 new biochorologic sub-provinces were proposed, providing unique palaeogeographic units for further studies of Early Devonian Gondwana. The application of biochorologic similarity to geological and palaeogeographical hypotheses is here shown to be a valuable tool or set of supplementary evidence. Illustrated herein, one of the criticisms of the use of biogeographical method has been answered to: biotic similarity is resultant of a continuous geology within Early Devonian Australasia. Within this method, faunal compositions are seen rather as a corollary of geological action. Our approach presents an attempt for biogeography to move away from narratives and towards testable palaeobiogeographic hypotheses. Appendix A. Supplementary data Supplementary data to this article can be found online at http://dx. doi.org/10.1016/j.palaeo.2015.11.037. References Adams, C.J., Korsch, R.J., Griffin, W.L., 2013. Provenance comparisons between the Nambucca Block, Eastern Australia and the Torlesse Composite Terrane, New Zealand: connections and implications from detrital zircon age patterns. Aust. J. Earth Sci. Int. Geosci. J. Geol. Soc. Aust. 60, 241–253. Afanasieva, M.S., Amon, E.O., 2013. Paleobiogeographical Radiolarian Provinces in the Devonian. Paleontol. J. 47, 1135–1154. Aldridge, R.J., Purnell, M.A., 1996. The conodont controversies. TREE 11, 463–468. Andreu, B., Colin, J.-P., Singh, J., 2007. Cretaceous (Albian to Coniacian) Ostracodes from the Subsurface of the Jaisalmer Basin. Micropaleontology 53, 345–370. Anstey, R.L., Pachut, J.F., Tuckey, M.E., 2003. Patterns of Bryozoan endemism through the Ordovician–Silurian transition. Paleobiology 29, 305–328. Arnold, G.O., Henderson, R.A., 1976. Lower palaeozoic history of the Southwestern broken river province, North Queensland. J. Geol. Soc. Aust. Int. Geosci. J. Geol. Soc. Aust. 23, 73–93. Berry, C., Giesen, P., Stein, W., Mannolini, F., VanAller Hernick, L., Landing, E., Wang, Y., 2009. Towards and Evaluation of the role of Mid Devonian Forests in the Devolopment of the Earth System. Geophys. Res. Abstr. 11. Boger, S.D., 2011. Antactica- Before and after Gondwana. Gondwana Res. 19, 335–371. Boucot, A.J., Scotese, C., Morley, R.J., 2013. Phanerozoic Paleoclimate: An Atlas of Lithologic Indicators of Climate. Society for Sedimentary Geology Concepts in Sedimentology and Paleontology 11, 1–478. Cawood, P.A., 2005. Terra Australis Orogen: Rodinia breakup and development of the Pacific and Iapetus margins of Gondwana during the Neoproterozoic and Paleozoic. Earth Sci. Rev. 69, 249–279. Cawood, P.A., Leitch, E.C., 1985. Accretion and dispersal tectonics of the Southern New England Fold Belt, Eastern Australia. In: Howell, D.G. (Ed.), Tectonostratigraphic terranes of the Circum-Pacific region. Circum-Pacific Council for Energy and Mineral Resources, Texas, pp. 481–492. Cayley, R.A., 2011. Exotic crustal block accretion to the eastern Gondwanaland margin in the Late Cambrian–Tasmania, the Selwyn Block, and implications for the Cambrian– Silurian evolution of the Ross, Delamerian, and Lachlan orogens. Gondwana Res. 19, 628–649. Cayley, R.A., Taylor, D.H., VandenBerg, A.H.M., Moore, D.H., 2002. Proterzoic–Early Palaeozoic rocks and the Tyennan Orogeny in central Victoria: the Selwyn Block and its tectonic implications. Aust. J. Earth Sci. 49, 225–254.
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The Early Devonian palaeobiogeography of Eastern Australasia Elizabeth M. Dowding ⁎, Malte C. Ebach School of Biological, Earth & Environmental Sciences, UNSW, Kensington, NSW 2052, Australia
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Article history: Received 27 August 2015 Received in revised form 6 November 2015 Accepted 24 November 2015 Available online 12 December 2015 Keywords: Australia Biochorema New Zealand Parsimony analysis of endemicity Tectonostratigraphic terranes
a b s t r a c t Biogeographical analyses of Devonian Australasia (Australia and New Zealand) have previously presented a narrative approach to the interrelationships of continental faunal provinces. Useful for the creation of biotic hypotheses, similarities and differences between biotic provinces remains untested. This study seeks to propose the first regional faunal provinces allowing a test of geological and biotic hypotheses using Parsimony Analysis of Endemicity. The relationships of eastern Australasian terranes were tested through the hierarchical analysis of faunal composition. Biochorologic sub-provinces were proposed using geo-spatial computer software and analysed through phylogenetic software that resulted in the formalisation of 20 biochorologic sub-provinces. The relationships between these units show a significant holdover of Silurian species, the interactions between convergent terranes and the continental margin, the distinct history of the Lachlan Fold Belt terranes, and offers biotic support for the existence of the Tasmanian microcontinent Vandieland. © 2015 Elsevier B.V. All rights reserved.
1. Introduction Encompassing 70% of the continental area, Devonian Gondwana saw one of the earliest major adaptive radiations in Earth's history, giving rise to many evolutionary important taxa (Scotese et al., 1999; Talent et al., 2000; Boucot et al., 2013). With a collection of tectonostratigraphic terranes (‘terranes’, fragmentary crustal segments sutured to a continental margin presenting unique and distinctive geological history), subduction zones, and back-arc and forearc basins, Gondwana's Early Devonian presented a dynamic environment and myriad habitats (Fig. 1, McElhinny et al., 2003). Proposed by Young (1981), Eastern Gondwana, encompassing Australasia, forms the Wuttagoonaspid–Phyllolepid faunal province. This province stands distinct from the remaining four provinces: the Cephalaspid province (Euramerica), Amphiaspid province (Siberia), Tannuaspid province (Tuva), and Galeaspid–Yunnanolepid (South China) (Young, 1981). Based primarily upon fish fauna, biogeographical studies describing continental relationships have also been made using brachiopods (Talent et al., 1972; Yolkin et al., 2000), conodonts (Talent et al., 1972), radiolarians (Afanasieva and Amon, 2013), gastropods (Cook, 2001), trilobites, sponges and polyzoans (Talent et al., 1972) respectively. Single taxa studies, however, inevitably leave informational gaps which prove problematic considering that associations and affinities drawn are clouded by the individual group's evolutionary history and relationships. Furthermore, for the creation of true representations of biotic areas, data from multiple taxa types, habits, and life histories ought to be incorporated; the patterns and conclusions drawn from ⁎ Corresponding author. E-mail address: [email protected] (E.M. Dowding).
http://dx.doi.org/10.1016/j.palaeo.2015.11.037 0031-0182/© 2015 Elsevier B.V. All rights reserved.
single taxa studies inherently missing significant palaeobiogeographical patterns (Damborenea and Macenido, 1992). A solution is to incorporate the distribution of multiple unrelated taxa in order to determine biotic overlap. Biochorologic units, such as biochorema and biochorologic sub-provinces, are endemic biotic centres within a geographic envelope alter in time, distribution, and scale, offering spatial and temporal analysis of endemism and distribution (Cecca and Westermann, 2003). Such an approach is seen to give the best coverage of the habited area, relieving in part absence signification due to the specimen's inability to survive geological action or fossilisation (Aldridge and Purnell, 1996; Girard and Renaud, 2011). For example, the coastline of the Early Devonian Australian margin is based upon the zonation of the conodonts Pedavis pesavis and Icriodus woschmidti (Mawson and Talent, 2003; Fig. 2). The ubiquitous nature of conodont deposits have seen them used as identifiers for biostratigraphy (Lyons and Percival, 2002; Mawson and Talent, 2003; Girard and Renaud, 2011). However, within the convoluted frame of a convergent back-arc basin, the use of a single taxon maintains its difficulties. Marine transgressions were a known feature of the Australian margin, supplemented by continuous marine deposition within a back-arc and the Wagga Marginal Sea, an ambiguity left unresolved by Talent et al. (2000) (Glen and Walshe, 1999; Glen, 2013). Whilst their palaeoecology makes no part of biochorologic characterisation, the formation of faunal sub-provinces will logically display the patterns of their deposition. The proposal of Talent et al. (2000) is contradicted by the presence of the marine genus of brachiopod Chonetes within deposits from the Wagga–Omeo terrane (VandenBerg et al., 1976). Endemic centres created through data gained from multiple taxa are an established approach for the hierarchical analysis of areas (Cecca and Westermann, 2003). These centres are bound by the
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Fig. 1. Devonian Gondwana, area of study highlighted. Reconstruction from Boger (2011).
temporary range limits of the constituent taxa, ranks scaling with the degree of endemism, taxa distribution, and duration (Westermann, 2000). The Early Devonian Australasian coast, presents a myriad of environments respectively affected and disrupted by terrane convergence and the shifting of the Gondwanan supercontinent (VandenBerg et al., 2000). During the Early Devonian, Gondwana was steadily moving towards Panthalassa, its active margin interacting with a geological collage of terranes and fragmentary blocks (Scotese et al., 1999; von Raumer and Stampfli, 2008; Fergusson, 2010; Boucot et al., 2013). Obduction, followed by subduction inversion formed the Early Devonian palaeotethys between the Gondwanan and Variscan terranes, East Gondwana proving convergent in a back-arc setting (Von Raumer et al., 2002; Fergusson, 2010; Boger, 2011). This movement, coupled with an alteration in the locus of convergence within Gondwana, set in motion a complex history of accretion, rifting, strike-slip faulting (Siluro-Devonian), and extensional/contractional phases across its eastern portion (Foster and Gray, 2000). During the course of this convergence, the geography and fauna are seen to have changed and altered with the geology (Lieberman, 2005). Silurian holdover faunas were preserved into the Early Devonian by the Wagga Marginal Sea and the rift and extensional basins that riddled the Australian coast (Talent et al., 1972). These Silurian fauna transitioned into more typical Devonian community compositions contemporaneously with the westward accretionary movements of the outboard terranes
(Veevers, 2004; YuanSheng et al., 2008). The nature of the relationships between these two coeval changes can be hierarchically represented, regional faunal provinces or ‘sub-provinces’ identified and their geological relationships inferred. Contrary to biochorologic and terrane models, tectonic models for Australasia are well established; the timing and nature of terrane accretions and basin formations being generally resolved within the Early Devonian (Cawood, 2005; Fergusson, 2010; Boger, 2011; Fergusson and Henderson, 2015). The Tasmanides are characterised by the Lachlan Fold Belt and Thomson Orogen, age of accretion younging from west to east (Foster and Gray, 2000; Glen et al., 2004). The central Lachlan Fold Belt, comprised of Howqua–Tabberabbera and Wagga-Omeo, exhibits an interesting correlation to the study of Yolkin et al. (2000). Analysing brachiopod community compositions, Yolkin et al. (2000) aligned the central Howqua–Tabberabbera and Molong–Monaro terranes of the eastern Lachlan Fold belt as a single biotic province depicting relation to Melbourne and New Zealand (Fig. 3). The constituent terranes of New Zealand were not delineated by Yolkin et al. (2000) and the relationships of these, consequently, will be sought here though the continuity of multiple taxa and bound by established terranes. The Melbourne, Tasmanian, and New Zealand terranes contain some of the most richly fossiliferous depositions within Australasia. Recently they have come under much scrutiny, their accretionary history generating many hypotheses such as ‘Vandieland’ (Cayley et al., 2002;
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Cayley, 2011), the reconstruction of Li and Powell (2001), and the placement of New Zealand between Australia and Antarctica (Ireland, 1992; Fagerstrom and Bradshaw, 2002). Described as a continental fragment, magmatic analysis of eastern Gondwanan granitoids have indicated that those of New Zealand, Northern Victoria Land, Marie Byrd Land in West Antarctica, Tasmania, and central Lachlan Fold belt (Howqua–Tabberabbera and Wagga-Omeo) were correlated, potentially representing a semicontinuous magmatic belt in excess of 2000 km along the Gondwanan margin, prior to subsequent rifting and breakup (Cooper and Millener, 1993; Daugherty et al., 1993; Muir et al., 1996). Evidence found within New Zealand's Reefton group, (the distribution and evolutionary history of endemic species), give biotic supplement to the established links between Tasmanian, Victorian, and Antarctic sedimentary basins (Fagerstrom and Bradshaw, 2002; Adams et al., 2013). The aim of this study is to propose the first Early Devonian biochorologic sub-provinces, presenting their hierarchical relationships to discern the affinities and geological relations of tectonostratigraphic terranes within Australasia. Focussing upon the Australasian coastline during the early Devonian (419–393mMa), this study will apply Parsimony Analysis of Endemicity (PAE) to existing hypotheses of terrane history (Young et al., 2010), palaeogeography (Talent et al., 2000) and biotic area evolution (Ebach and Edgecombe, 2001). Using faunal and biochorologic data, new regional configurations of Australasian geological and biotic associations will be proposed. This paper will also provide supplementary evidence for the existence of Vandieland, the location of New Zealand, and establish the nature of the relationship between the terranes of the Lachlan Fold Belt. Devonian biogeographical reconstructions have largely focussed upon information from single taxa (e.g. genera and families). The patterns and conclusions are drawn from these analyses focusing upon the life histories of single taxa and inherently miss significant palaeobiogeographical patterns (Parenti and Ebach, 2009). Similarly, geographical or geological approaches to reconstruction remain limited in their ability to suggest faunal distribution and seaboard. This study seeks to understand the dualism between the evolution of location and life. 2. Materials and Methods 2.1. Biogeographic Setting Fig. 2. The Early Devonian marine seaboard of Australia proposed by Talent et al. (2000).
Early Devonian (419–393 Ma) Australasia encompassed Australia and New Zealand respectively, New Guinea not yet having been formed.
Fig. 3. Faunal relationships based upon brachiopods of Early Devonian crustal blocks (Yolkin et al., 2000).
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Located along the Gondwanan eastern coastline, Australasia exhibits 14 early Devonian tectonostratigraphic terranes and basins, the former being fault bound regions with a unique stratigraphic history (Leitch and Scheibner, 1987). These units are generally allochthonous and present dynamic histories of rift and suture, interacting with a convergent plate boundary (Cawood and Leitch, 1985). The 14 Devonian Australasian terranes and basins are used for analysis of geological evolution based upon their faunal characteristics. The relationships between these terranes will be assessed through the compositions of their biota. Selected terranes are chosen based on endemic taxa, in particular trilobites, brachiopods, fish, sarcopterygians, and placoderm faunas. The terranes were required to show some overlapping faunal distributions, and have been verging upon or accreted to the Gondwanan margin after the Early Ordovician and before the Early Devonian. 2.2. Data Collection The following data was collected manually through an extensive review of the existing literature, such as, the terrane maps of Leitch and Scheibner (1987) and Vandenberg et al. (2000) for Australia and New Zealand (see Appendix A, Table 1). The data includes all recorded brachiopods, trilobites, sarcopterygians, and placoderms from 14 terranes and basins. 91 Early Devonian (419–393 Ma) formations across 14 terranes and basins were described in terms of type, composition, and environment of deposition. The tectonostratigraphic terranes, being fragmentary, unique, lithological units, were required to have both endemic and more cosmopolitan inhabitant biota (VandenBerg et al., 2000). Devonian corals and conodonts were discounted due to their ubiquitous distributions in favour of brachiopods, trilobites, sarcopterygians, and placoderms. Patterns of endemicity and distribution would be clouded by the informational bias presented by the cosmopolitan nature of these taxa (Young, 1987). Accordingly, sarcopterygian, placoderm, trilobite, and brachiopod presences were recorded within formation and corollary terrane, data pulled from previous palaeontological works (Table 1). Genus, rather than species, was retained in order to account for disarticulation and the inability to specifically identify all specimens within fossil record, allowing a display of higher order relationships and patterns (Upchurch et al., 2002). The location of the specimen was noted in terms of latitude and longitude, and when the specific location was unavailable, the formations location was recorded. Taxon age was not incorporated to analysis; distribution, composition, and location used for character individuation (Young, 1987, 2007). 2.3. Terranes and basins The following terranes and basins were used in the PAE analysis: Buller Terrane. — Part of Devonian New Zealand the Buller Terrane comprises 11 fossiliferous formations. The formations of this terrane exhibit a high energy, near shore beach to shelf depositional environment. Canning Basin. — Used as an outgroup to root the tree. No formations were noted for the intercratonic basin, with low energy, deep-water to hypabyssal deposition. Darling Basin. — This basin was muddy, slowly subsiding, and asymmetrical; a shallow marine depositional environment with deltaic, fluvial sediments. Georgina Basin. — Cratonic Australian segment presenting two fossiliferous Devonian formations. Non-marine, low energy basinal palaeoenvironment. Hodgkinson–Camel Creek Terrane. — Fossils were found in a medium to low energy ocean floor and trench accretionary prism.
Howqua–Tabberabera Terrane. — This composite terrane is comprised of 4 individual formations and the Wentworth group. Howqua–Tabberabera exhibited a muddy bottomed, lacustrine to shallow marine depositional environment. Bioherms have been noted within the terrane, suggesting medium to low energy environments. Melbourne Terrane. — 17 formations are found to represent deep water palaeoenvironment. Medium to low energy environment can be seen through the transport channels and rippling that marks the terranes beds, laminations, and formations. Located at the back of the Kanmantoo orogeny, it was a foreland basin moving into orogenic deformation during the Middle Devonian Tabberabberan orogeny. Molong–Monaro Terrane. — This composite terrane exhibits traits of a shallow marine shelf. The 27 formations vary between low energy muddy substrates, volcaniclastics, and patch reefs complete with reef mounds and some megaturbidites forming within the forearc basin. Molong–Monaro presents basins and rifting, both of which were associated with terrane dispersal. The tectonic highs, between the troughs and basins, were progressively closed during the Devonian by the twin actions of the Tabberabberan and Bowning orogenic events. Narooma Terrane. — The Devonian segment of this allocthonous terrane is limited to the Bunga Beds. Situated against an escarpment within a volcanic graben, the Bunga Beds present deep water deposition, a freshwater lake dammed by the graben. The Narooma Terrane was accreted during the Benambran orogeny, presenting outer arc slope and trench during the early Devonian (Stokes et al., 2015). Sydney–Bowen Basin. — Volcanically active, the Sydney–Bowen Basin presents shallow marine and terrestrial sedimentation, succeeded by deeper water. High energy conditions gradually decrease with the increasing water depths into a foredeep. Tamworth–Yarrol Terrane. — This composite terrane (Wisemans Arms, Woolomin, Cockburn, and Texas terranes) encompasses a forearc basin in the Graveyard Creek foredeep and the Kangaroo Hills deep, with two superimposed arc fringe-forearc basin sequences. Extensively bioturbated, this terrane had a low energy environment characterised by siltstone and sandstone and some volcaniclastics with shallow water environments in the west found within six formations. North-West Tasmanian Terrane. — Though a low to medium energy environment, there is evidence of strong current activity within this terrane. Rapid deposition without much washing can be seen in this uplifting depositional interface which moved into shallow water. This pattern can be seen graded within each of the seven North-West Tasmanian formations. Takaka Terrane. — Steep eastward dipping has been noted within the Takaka terrane. Characterised by the Devonian Baton formation, Takaka is a mud rich, deep offshore environment with present turbidites. Wagga-Omeo Terrane. — This relatively low energy terrane presents a shallow marine depositional environment as seen across its 6 Devonian formations. During the Devonian the Wagga-Omeo Terrane underwent dispersal of the order of kilometres to tens of kilometres, rifting throughout the Early Devonian and Late Silurian.
2.4. Biochorologic sub-provinces Biochorema are endemic biotic centres within a geographic envelope and therefore alter in time, distribution, and scale (Cecca and Westermann, 2003). For this regional study, biochorema are referred
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to as sub-provinces. 20 biochorologic sub-provinces are proposed, characterised by 10% endemicity of fossil genera; the recommended unit for provincial studies (Guangrong, 1998; Anstey et al., 2003; Andreu et al., 2007). These biochorologic sub-provinces were identified using Biodiverse 0.19, a spatial analysis tool for the analysis of diversity, which distinctively offers the ability to map and analyse georeferenced biological specimens (Laffan et al., 2010). Within this analysis, clustered taxonomic dissimilarity relationships were used and areas and taxa that did not achieve 10% endemicity were removed. In this way, Biodiverse allows a distinctive insight to the similarity and dissimilarity of spatial diversity, displaying endemic centres and presenting relationships within a hierarchical phenogram. The recognition of a hierarchy within the results leads to the implication, following from Anstey et al. (2003), that smaller units may be nested within larger ones. The majority of biogeographical studies (Young, 1981; Rich and Young, 1996; Young et al., 2010) have presented large geographic areas, this study seeking to unbind the nested relationships seen within continental studies. This recognition of biogeographical hierarchy allows the discernment of how endemic regions or sub-provinces have been formed through ascertainable evolutionary process (Rosen, 1988; Rosen and Smith, 1988; Anstey et al., 2003). Because of the proximity of each of the deposits, differences seen between the sub-provinces are assumed to be related more to geological action and consequent habitat type as opposed to climatic influence (Yanin, 2010). To this end, polytypic areas prove especially advantageous, giving equal valency to all habitat types and capturing signals given by mixed or transitional zones and environments (Guangrong, 1998).
2.5. Analysis of tectonostratigraphic terranes Tectonostratigraphic terranes, being fragmentary crustal segments sutured to a continental margin presenting unique and distinctive geological history, were analysed by their biotic similarities. Proposed by Rosen (1988) and Rosen and Smith (1988) endemicity data, hierarchically structured within a parsimonious phenogram, can be used as a pluralistic approach to discerning geological and stratigraphic relations. The use of biotic data for area classifications has been widely utilised following Rosen's (1988) Parsimony Analysis of Endemicity (PAE). PAE defines endemic areas, using these to determine hypothetical relationships in the absence of phylogenetic data (Morrone and Escalante, 2002; Nihei, 2006; Morrone, 2014a). Based upon taxa presence-absence data matrices, PAE constructs parsimonious cladograms allowing the identification of ‘biotic components’: historical biotas, biogeographical assemblages, taxonomic assemblages, species assemblages, and their hypothetical relationships (Garcia-Barros et al., 2002; Morrone, 2014b). For each terrane, the genera profile was used to provide character individuation; distributional data entered as a presence ‘1’ only matrix and analysed using the parenthetical cladistics program LisBeth v.1.3 (Zaragueta Bagils et al., 2012). Noted within critiques, PAE groups on absences, thereby creating similarities between areas that share no taxa. Many have argued that this fails to capture historical complexity (Morrone and Escalante, 2002; Garzon-Orduna et al., 2008). Binary forms, such as PAE, often become problematic in the face of an incomplete fossil record, as relationships based upon sparse or well distributed data are at risk of losing presence significance to overwhelming sequenced absence data (absence signification). The use of LisBeth v. 1.3 within the present study, applies a parenthetical matrix, thereby allowing only presences. LisBeth v.1y3 was run using three item analysis and compatibility (3ia function), finding the most optimal (i.e. compatible) tree from the largest sum of fractional weighting (Nelson and Ladiges, 1996). Characters are depicted as hierarchies or polarized and ordered tree states, congruent or optimal tress summarised within the intersection tree (Zaragueta Bagils et al., 2012).
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Following from Young (1987), temporal information was seen to be an incorporated feature of the taxon or terrane, considered at terminal braches rather than nodes. This option is available due to the omission of phylogenetic data (Upchurch and Hunn, 2002), namely that distributions seen as indicative of geology rather than the specific evolutionary history of a taxon. 3. Results The apical tectonostratigraphic terranes (Fig. 4) correlate to the allochthonous VandieLand of Cayley (2011). The New Zealand terranes, Takaka and Buller, are proposed to have interacted prior to the accretion of Vandieland during the late Cambrian–Ordovician (Cooper and Millener, 1993). The central segment of the terrane tree intimates detail of the interactions of the Macquarie Arc and Narooma Accretionary Complex, assumedly falling central due to their shared history with the Howqua–Tabberabbera and Wagga-Omeo terranes (Table 1). The terrestrial, freshwater, mixed profile basins and terranes of Hodgkinson– Camel Creek and the Georgina Basin, basally group. Through the Biodiverse analysis of spatial cluster and endemicity, 20 biochorologic sub-provinces are proposed across the region, distribution allowing the proposed configuration of the Australian seaboard (Fig. 5), Molong-Monaro and Melbourne terranes offer the highest number of sub-provinces across time (Table 2). The analysis of the biochorologic subregions (Fig. 6) is seen to mirror those of the terranes (Fig. 4). The apex of the tree (Fig. 6) broadly reflects the terrane relationships of Fig. 4, grouping the endemic centres that fell within the Melbourne, Molong–Monaro, Buller, North-West Tasmanian, Howqua– Tabberabbera terranes, and the Sydney–Bowen Basin. Interestingly, the base of the tree indicates a temporal split, Pragian and Emsian faunal distributions grouping together, Lochkovian sub-provinces dominating the upper half of the tree. 4. Discussion Spatially and temporally bracketed 20 new biochorologic subprovinces give crucial insight to the biotic development of Eastern Australasia, its palaeogeography, and the dependence upon geological action through time (Fig. 5, Appendix A, Table 2). The interrelationships of these biochorologic units to their facilitating geology offer supplement to
Fig. 4. Analysis of terranes. The analysis of the terrane data returned one tree with a completeness (CL) of 92.0% and a retention index (RI) of 0.76.
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Fig. 5. a. Seaboard for the Early Devonian of Australia following the marine line of the Sydney–Bowen Basin and the biochorologic subprovinces. 1 — Mulga (Darling Basin), 2 — Dulcie (Georgina Basin), 3 — Cravens-Toomba (Georgina Basin), 4 — Ukalunda (Sydney–Bowen Basin), 5 — Humevale (Melbourne), 6 — Amphitheatre (Darling Basin), 7 — Boola (Melbourne), 8 — Florence (North-West Tasmania), 9 — Trundle (Wagga-Omeo), 10 — Walhalla-Kilgower (Melbourne and Howqua–Tabberabbera), 11 — Derriwong (Wagga-Omeo), 12 — Mandagery (Molong–Monaro), 13 — Cavan (Molong–Monaro), 14 — Garra (Molong–Monaro), 15 — Crudine (Molong–Monaro), 16 — Bowning (Molong–Monaro), 17 — Boyd (Narooma), 18 — Wurutwun (Melbourne), b. New Zealand.19 — Reefton (Buller), 20 — Baton (Takaka).
geologic models, providing evidence of the rotation of the Victorian region and Melbourne zone, the closure of the Wagga Marginal Sea, the nature of the relationship of continental basins and allochthonous terranes, and the biotic affinities of the three segments of the Lachlan Fold Belt. The analogous nature of the relationship between geologically based models and the results seen here confirm the efficacy of our palaeobiogeographic approach to regional biotic and geologic hypotheses. 4.1. Evidence for interaction between the Darling Basin the Lachlan and Thomson Orogens The Darling Basin covers two Biochorologic sub-provinces, being the Emsian Mulga (Table 2, number 1) and the Lochkovian to Pragian Amphitheatre (Table 2, number 6). Previously given only perfunctory attention (Fergusson, 2010; Glen et al., 2013), the results have indicated
the Darling Basin was a dynamic region, influenced by the movements of the Lachlan and Thomson Orogens making it both a cosmopolitan hub (Amphitheatre sub-province) and subsequently a refuge basin (Mulga sub-province) (Fig. 6). The analysis indicates that the Amphitheatre sub-province had biotic exchange with the sub-provinces of the Wagga-Omeo terrane. During the Lochkovian, the Amphitheatre sub-province was flanking the Wagga Volcanic Arch, overlap sequences of the Cobar-supergroup deposited within the rifts and flanking shelves of both the Lachlan and Thompson Orogens (Webby, 1972; Glen et al., 1994; Fergusson, 2010; Glen, 2013). Initially afforded by a period of major extension within the region, the relationship between the Darling Basin and the Orogens is limited to the Lochkovian. Due to interplay between the orogens, the Cobar basin was obliquely opened and then closed forcing the cessation of the eastward flowing
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problematic reconstructions (Fig. 2). During the early Devonian the Wagga Marginal Sea was closed by the continuous rollback of the subduction hinge coupled with regional uplift and strike-slip faulting (Lambeck and Stephenson, 1986; Fergusson, 2010). These movements lead to increasing endemism amongst the LFB subprovinces, characterised by refuge basins (Crook et al., 1973; Crook, 1980; Strusz, 2000), and intermediate and freshwater environments. The taxa profile have drawn the seaboard for the Pragian (Fig. 5), the Wagga marginal sea shown to not be fully closed, limited by the mixed profile of the Cavan sub-province (Table 2, number 13). The depiction of the LFB as a single geological unit was to an extent mirrored in Yolkin et al. (2000), who used brachiopod genera to group the Lachlan Fold Belt terranes of Howqua–Tabberabbera and Molong– Monaro into a single biotic unit. This study, however, has shown that Howqua–Tabberabbera and Molong–Monaro not only have distinct biochorologic sub-provinces (Table 2), but also that these subprovinces are more closely related to those of the Melbourne terrane than either are to each other (Fig. 6). Furthermore, during the early Devonian the Wagga margin was subducting between Gondwana and the Molong Island arc, maintaining the separation of the Howqua– Tabberabbera and Molong Monaro terranes (Fergusson, 2010). Fig. 6. The analysis of biochorlogic sub-provinces. The tree presents a completeness of 75.3% and a retention index (RI) of 0.808.
palaeocurrent (Glen and Walshe, 1999). The termination of this current, coupled with the closure of the Wagga Marginal Sea lead to the isolation of Darling Basin, the fauna of the Amphitheatre sub-province transitioning into those of the Mulga. The Mulga sub-province is proposed here to have been a refuge basin, the persistence of late Silurian and Lochkovian fauna into to the Emsian apparent through the close relationship of the Darling Basin sub-provinces to each other than to another area (Table 2, Fig. 6). 4.2. The shared biochorologic sub-province of Hodgkinson–Camel Creek and Georgina Basin The Hodgkinson–Camel Creek terrane is one of Australia's youngest accretionary complexes (Withnall and Henderson, 2012). Formed within a Siluro-Devonian forearc accretionary prism the Cravens-Tooma sub-province (Table 2, number 3) exhibits the characteristics of a convergent margin with widespread, deep marine turbidites. The subprovince is, however, found to have been equally influenced by the continental margin (De Keyser and Lucas, 1968; Arnold and Henderson, 1976; Day et al., 1978). Constructed jointly out of the convergent margin and the continental freshwater of the Georgina basin, CravensToomba represents a composite sub-province (Table 2). Terrigenous materials and freshwater species were deposited into the marine Hodgkinson basin by strong, heavily laden currents from the west which lost most of their load and force as it moved towards the more distal parts of the basin (de Keyser and Lucas, 1968; Haines et al., 2001; Li and Powell, 2001). Whilst the exact linkages between the inland Georgina Basin and the convergent Hodgkinson–Camel Creek terrane cannot be directly ascertained, this sub-province does provide crucial insight into the interactions between inboard terranes and the Gondwanan margin. 4.3. The discrete history of the Lachlan Fold Belt terranes Often depicted as a single accretionary unit (Li and Powell, 2001; Boger, 2011), the terranes of the Lachlan Fold Belt exhibits a suite of unique biotic sub-provinces (Tab 2). These relationships between the sub-provinces of the Lachlan Fold Belt are often contained within the borders of their host terrane, biotic similarities offering insight into the southern extension of the LFB, a zone which has been prone to
4.4. Vandieland The Melbourne terrane exhibits four sub-provinces including Humevale, Boola, Walhalla-Kilgower, and Wurutwun (Table 2 number 7, 10, and 18). During the early Devonian the Melbourne zone formed a depositional trough, sediments from the Stawell, and Bendigo zones, Selwyn Block inclusive, providing sedimentary material (Fergusson, 2003; Cayley, 2011). This external influence upon the terranes subprovinces coupled within the strong affiliations between them and those of North West Tasmanian provide biotic evidence for Vandieland (Cayley et al., 2002; Cayley, 2011). Vandieland, composed of the Selwyn block, the Melbourne Zone, and Tasmania was proposed by Cayley et al. (2002) and Cayley (2011) to explain the trapped pre-Cambrian continental extension of Tasmania, being the Selwyn block, and its occurrence upon mainland Australia (Crawford et al., 2007; Berry et al., 2009; Moore et al., 2015). Rotation of the Melbourne zone, evidenced by the migrating axis of trough deposition, which moved eastward providing links to the late early Devonian Molong-Monaro sub-province Cavan, and influencing the closure of Wagga Marginal Sea (Webby, 1972; Lambeck and Stephenson, 1986; Miller and Gray, 1996; Cayley et al., 2002). The delineation of this microcontinent and its allochthonous history is understood to provide a unifying framework for the explanation of the ‘Selwyn paradox’, and the observed geological connections between Tasmania, New Victoria Land, and Victoria (Veevers, 2004; Cayley, 2011; Moore et al., 2015). 4.5. New Zealand Strong association between Baton, Reefton, Humevale, and Florence suggest a proximal relationship between NZ and the southern Australian margin (Fig. 6). Previous studies have indicated that New Zealand was located between the 30th and 40th palaeolatitidue (Muir et al., 1996; Fagerstrom and Bradshaw, 2002; Torsvick and Cocks, 2013), a location which would allow exchange between Australian populations and those previously observed within Antarctic sedimentary basins (Adams et al., 2013). Magmatic analysis of granitoids have indicated that those of New Zealand, Northern Victoria Land, Marie Byrd Land in West Antarctica, Tasmania, and the central Lachlan Fold belt (Howqua -Tabberabbera and Wagga-Omeo) were correlated, potentially representing a semicontinuous magmatic belt in excess of 2000 km along the Gondwanan margin (Cooper, 1989; Cooper and Tulloch, 1992; Daugherty et al., 1993; Muir et al., 1996). Tectonically activated variation, with particular regard to the extensional and contractional
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phases of the LFB, altered the nature of both sedimentation and habitat, consequently shifting affiliations of the Lachlan Fold Belt away from the uninterruptedly stable sub-provinces of Tasmania and New Zealand (Fig. 6). The placement of New Zealand between the 30th and 40th latitude is therefore seen to complement the temporal changes marked between these New Zealand sub-provinces and the Victorian terranes. Both the New Zealand and Tasmanian terranes were primarily unaffected by the contractional, extensional phases of the Lachlan Fold Belt and consequently their links to the aforementioned remain strongest within the earliest Devonian (Fig. 6). 5. Conclusion Devonian Gondwana saw one of the earliest major adaptive radiations in Earth's history, giving rise to many evolutionarily important taxa (Scotese et al., 1999; Talent et al., 2000). This study presents the first regional palaeobiogeographic analysis of Early Devonian Australasia across a variety of taxonomic groups. Through the use of these multiple taxa, 20 new biochorologic sub-provinces were proposed, providing unique palaeogeographic units for further studies of Early Devonian Gondwana. The application of biochorologic similarity to geological and palaeogeographical hypotheses is here shown to be a valuable tool or set of supplementary evidence. Illustrated herein, one of the criticisms of the use of biogeographical method has been answered to: biotic similarity is resultant of a continuous geology within Early Devonian Australasia. Within this method, faunal compositions are seen rather as a corollary of geological action. Our approach presents an attempt for biogeography to move away from narratives and towards testable palaeobiogeographic hypotheses. Appendix A. Supplementary data Supplementary data to this article can be found online at http://dx. doi.org/10.1016/j.palaeo.2015.11.037. References Adams, C.J., Korsch, R.J., Griffin, W.L., 2013. Provenance comparisons between the Nambucca Block, Eastern Australia and the Torlesse Composite Terrane, New Zealand: connections and implications from detrital zircon age patterns. Aust. J. Earth Sci. Int. Geosci. J. Geol. Soc. Aust. 60, 241–253. Afanasieva, M.S., Amon, E.O., 2013. Paleobiogeographical Radiolarian Provinces in the Devonian. Paleontol. J. 47, 1135–1154. Aldridge, R.J., Purnell, M.A., 1996. The conodont controversies. TREE 11, 463–468. Andreu, B., Colin, J.-P., Singh, J., 2007. Cretaceous (Albian to Coniacian) Ostracodes from the Subsurface of the Jaisalmer Basin. Micropaleontology 53, 345–370. Anstey, R.L., Pachut, J.F., Tuckey, M.E., 2003. Patterns of Bryozoan endemism through the Ordovician–Silurian transition. Paleobiology 29, 305–328. Arnold, G.O., Henderson, R.A., 1976. Lower palaeozoic history of the Southwestern broken river province, North Queensland. J. Geol. Soc. Aust. Int. Geosci. J. Geol. Soc. Aust. 23, 73–93. Berry, C., Giesen, P., Stein, W., Mannolini, F., VanAller Hernick, L., Landing, E., Wang, Y., 2009. Towards and Evaluation of the role of Mid Devonian Forests in the Devolopment of the Earth System. Geophys. Res. Abstr. 11. Boger, S.D., 2011. Antactica- Before and after Gondwana. Gondwana Res. 19, 335–371. Boucot, A.J., Scotese, C., Morley, R.J., 2013. Phanerozoic Paleoclimate: An Atlas of Lithologic Indicators of Climate. Society for Sedimentary Geology Concepts in Sedimentology and Paleontology 11, 1–478. Cawood, P.A., 2005. Terra Australis Orogen: Rodinia breakup and development of the Pacific and Iapetus margins of Gondwana during the Neoproterozoic and Paleozoic. Earth Sci. Rev. 69, 249–279. Cawood, P.A., Leitch, E.C., 1985. Accretion and dispersal tectonics of the Southern New England Fold Belt, Eastern Australia. In: Howell, D.G. (Ed.), Tectonostratigraphic terranes of the Circum-Pacific region. Circum-Pacific Council for Energy and Mineral Resources, Texas, pp. 481–492. Cayley, R.A., 2011. Exotic crustal block accretion to the eastern Gondwanaland margin in the Late Cambrian–Tasmania, the Selwyn Block, and implications for the Cambrian– Silurian evolution of the Ross, Delamerian, and Lachlan orogens. Gondwana Res. 19, 628–649. Cayley, R.A., Taylor, D.H., VandenBerg, A.H.M., Moore, D.H., 2002. Proterzoic–Early Palaeozoic rocks and the Tyennan Orogeny in central Victoria: the Selwyn Block and its tectonic implications. Aust. J. Earth Sci. 49, 225–254.
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