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Bryant, Litticia& Krosch, Matthew (2016) Lines in the land: a review of evidence for eastern Australia’s major bio- geographical barriers to closed forest taxa. Biological Journal of the Linnean Society, 119(2), pp. 238-264.

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Notice: Please note that this document may not be the Version of Record (i.e. published version) of the work. Author manuscript versions (as Sub- mitted for peer review or as Accepted for publication after peer review) can be identified by an absence of publisher branding and/or typeset appear- ance. If there is any doubt, please refer to the published source. https://doi.org/10.1111/bij.12821 1 Title: Lines in the land: a review of evidence for eastern Australia’s major biogeographical barriers to 2 closed forest taxa. 3 4 Short title: Eastern Australian biogeographical barriers 5 6 Bryant, L.M.1, Krosch, M.N.1,* 7 8 1 School of Earth, Environmental and Biological Sciences, Queensland University of Technology, 9 Brisbane 4000, Australia. 10 11 12 * Corresponding author. Email: [email protected] 13 14 15 Keywords: fragmentation, refugia, Pleistocene, Miocene, divergence, aridification 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 Abstract 53 54 The influence of climatic changes occurring since the late Miocene on Australia’s eastern mesic 55 ecosystems has received significant attention over the past twenty years. In particular, the impact of 56 the dramatic shift from widespread rainforest habitat to a much drier landscape in which closed 57 forest refugia were dissected by open woodland/savannah ecosystems has long been a focal point in 58 Australian ecology and biogeography. Several specific regions along the eastern coast have been 59 identified previously as potentially representing major biogeographical disjunctions for closed forest 60 taxa. Initially, evidence stemmed from recognition of common zones where avian species/ 61 subspecies distributions and/or floral communities were consistently separated, but the body of 62 work has since grown significantly with the rise of molecular phylogeographic tools and there is now 63 a significant literature base that discusses the drivers, processes and effects of these hypothesised 64 major biogeographical junctions (termed barriers). Here, we review the literature concerning eight 65 major barriers argued to have influenced closed forest taxa; namely, the Laura Basin, Black 66 Mountain Corridor, Burdekin Gap, Saint Lawrence Gap, Brisbane Valley Barrier, Hunter Valley 67 Barrier, Southern Transition Zone and East Gippsland Barrier. We synthesise reported 68 phylogeographical patterns and the inferred timing of influence with current climatic, vegetation 69 and geological characteristics for each barrier to provide insights into regional evolution and seek to 70 elicit common trends. All eight putative biogeographical barriers are characterised currently by 71 lowland zones of drier, warmer, more open woodland and savannah habitat, with adjacent closed 72 forest habitats isolated to upland cool, wet refugia. Molecular divergence estimates suggest two 73 pulses of divergence, one in the early Miocene (~20‐15Mya) and a later one from the Pliocene‐ 74 Pleistocene (~6‐0.04Mya). We conclude with a prospectus for future research on the eastern 75 Australian closed forests and highlight critical issues for ongoing studies of biogeographical barriers 76 worldwide. 77 78 Introduction 79 80 The concept that habitat fragmentation can drive divergence and speciation of isolated populations 81 is fundamentally enshrined in the modern ecological frameworks of island biogeography, equilibrium 82 theory and species turnover (MacArthur & Wilson, 1967). Vicariance or contraction of large 83 continuous habitats into one or more smaller habitat patches (refugia) that are isolated from any 84 other such patches by unsuitable territory across which taxa may not disperse (barriers) can 85 interrupt gene flow among populations, potentially leading to inbreeding, divergence in allopatry or 86 even extinction. By this definition, the two biogeographical phenomena are intrinsically interlinked, 87 as the distributions of taxa that are restricted to a refugium are necessarily circumscribed by a 88 barrier that they cannot cross. However, situations may also arise in which refugia may harbour 89 species of varying dispersal abilities and ecologies, with some unable to cross barriers and so remain 90 isolated, whilst others can move between refugia across zones of otherwise unsuitable habitat. This 91 scenario leads to the ‘leaky barrier’ concept, in which some taxa exhibit strong 92 phylogeographical/phylogenetic structuring across a barrier, whereas other co‐located sympatric 93 taxa show less, or no structuring (Avise 2000). Nonetheless, both refugia and barriers can be 94 considered equally important in determining and driving distributions and genetic structure of 95 isolated populations. Here we review published phylogeographical patterns across several putative 96 biogeographical barriers in eastern Australia to assess common trends and discordances among taxa. 97 98 For most of the Cretaceous‐Paleogene (145‐23 million years ago, Mya) eastern Australia possessed a 99 relatively continuous tract of mesic habitat that extended the entire length of the east coast and 100 deep into the interior of the continent (Sluiter & Kershaw, 1982; Truswell, 1993; Hill, 2004; Martin, 101 2006; Byrne et al., 2011). Regional climatic warming commencing during the Cenozoic (~65Mya) was 102 reinforced by Australia’s northward drift, and offset global cooling driven by the opening of the 103 Drake Passage ~35Mya (Barker & Burrell, 1977; Crook, 1981). Associated changes to ocean currents 104 and trade winds altered seasonal rainfall across the Australian mainland creating widespread 105 aridification (Wilford & Brown, 1994; Hill, 2004; Cook & Crisp, 2005; Byrne et al., 2011). Aridification 106 continued during the Miocene‐Pliocene epochs, driving reduction in closed forest by expansion of 107 dry, open sclerophyll woodland and grassland (Martin, 1982, 1990, 1998; VanDerWal, Shoo & 108 Williams, 2009; Byrne et al., 2011). Pollen records from northeast Queensland show a shift in floral 109 assemblages coincident with climatic fluctuations during these periods, with a much greater 110 occurrence of sclerophyllous taxa during dry periods (Hekel, 1972; Kershaw, 1976, 1994; MacPhail, 111 1997; Moss & Kershaw, 2000; Genever et al., 2003). Australia’s southern temperate rainforests also 112 underwent extensive contraction and vicariance since the mid‐Miocene (Martin, 1998; Crisp, Cook & 113 Steane, 2004; VanDerWal et al., 2009). Retreat of closed forest habitats into isolated, often upland, 114 refugia continued during the Pleistocene, particularly around 500 thousand years ago (kya) (Martin, 115 2006), and the Last Glacial Maximum (e.g., Kershaw, 1985, 1994; Nix & Switzer, 1991; Shulmeister & 116 Lees, 1995). The current interglacial period has been characterised by re‐expansion of closed forest 117 from many refugia across eastern Australia, and long‐term areas of persistence are inferred for the 118 Wet Tropics of northeast Queensland, and the Clarke, Border and McPherson Ranges of mid‐ and 119 southeast Queensland, respectively (Rosauer et al., 2015). 120 121 Vicariance of Australia’s closed forests thus resulted in an archipelago of refugia along the east 122 coast, separated by sometimes vast areas of drier open woodland/grassland. Dissecting this mosaic 123 of closed and open forests are eight proposed major biogeographical barriers ‐ areas of unsuitable 124 habitat that interrupt gene flow between populations isolated on either side (hereafter termed 125 simply ‘barriers’). These regions represent noted biotic transition zones and are invoked in 126 influencing the distribution, divergence and population structure of many closed forest‐restricted 127 groups (Byrne et al., 2011; Chapple et al., 2011a). These barriers include: the Laura Basin, Black 128 Mountain Corridor, Burdekin Gap, Saint Lawrence Gap, Brisbane Valley Barrier, Hunter Valley 129 Barrier, Southern Transition Zone, and East Gippsland Basin (Figure 1). These barriers delimit not just 130 refugial populations, but also form the boundaries of distinct phytogeographical regions/subregions 131 (Ebach et al. 2013; 2015, Figure 1). All of these subregions contain some remnant closed forest, and 132 many are noted hotspots of diversity, endemism and long‐term stability (Rosauer et al,. 2015). 133 134 The interaction between isolated refugial populations of closed forest‐restricted taxa and the 135 intervening dry habitat in eastern Australia has created an ideal model system for investigating the 136 role that vicariance has played in speciation and diversification (Moritz et al., 2000; Graham et al., 137 2006; Byrne et al. 2011; Chapple et al., 2011a). In this review, we collate, synthesise and critically 138 evaluate existing phylogeographical, palaeoecological, geological, climatic and vegetation 139 information for each of the major barriers to explore common trends and discuss discordances 140 among taxa in both pattern and timing of divergence. Typically, the general expectation for closed 141 forest taxa is that these barriers interrupted gene flow among bisected refugia, driving divergence of 142 disjunct populations. Divergence may be manifest at multiple evolutionary scales depending on the 143 organism in question: perhaps evident only in haplotype/allele frequency differences, or substantial 144 enough that separated populations may be considered different biological species with distributions 145 delimited by a known barrier. Further, estimated ages for molecular divergence events across 146 putative barriers should cluster around predicted ages (late Miocene‐Pleistocene, 10‐0.1Mya) for 147 closed forest contraction inferred from palaeovegetation data. Thus, single vicariance events for 148 each barrier represent the predominant hypotheses tested by studies reviewed here. By assimilating 149 published literature on these barriers into a ‘one‐stop shop’, we have identified some inconsistency 150 in the literature, as well as common trends in phylogeographical patterns among the putative 151 barriers, key knowledge gaps and avenues for potential future research. This review should be a 152 useful resource for phylogeographers working in Australia’s diverse mesic forest archipelagos and 153 encourage further work on this biogeographically dynamic region. 154 155 Laura Basin 156 157 Cape York Peninsula in north‐eastern Queensland boasts a complex underlying geology that 158 supports a wide variety of vegetation types. Eucalyptus woodlands dominate, but grasslands, 159 Melaleuca forest, mixed shrublands, mangroves and rainforest are well‐represented (Cofinas & 160 Creighton, 2001). Rainforest occurs mostly in the headwaters of the Jardine River at the very 161 northern tip of the Cape and, further south, in the higher elevation Iron and McIlwraith Ranges on 162 the eastern Cape. These large tracts of rainforest are separated from smaller high elevation 163 Eucalyptus‐dominated closed forests at Cape Melville by the geological feature known as the Laura 164 Basin, of which the Laura Lowlands bioregion is the surface product (Figure 2a,b) (Wellman, 1995). 165 The Laura Basin (LB) largely separates Cape Melville from the Wet Tropics south of Cooktown 166 excepting a narrow strip along the eastern coastline that possesses few small, poorly characterised, 167 closed forest isolates (e.g., Cape Flattery, Starcke Range – Figure 2a). The LB comprises a region of 168 ancient alluvial lowlands that is dominated by extensive savannah and open forest and encompasses 169 several river catchments (Figure 2b). These lowlands demarcate the northern limit of the Great 170 Dividing Range (GDR) and isolate the higher‐elevation Cape Melville from the McIlwraith Range to 171 the northwest and the GDR to the southeast. 172 173 The vegetation history of the LB is poorly known, and interpretations of regional patterns rely on 174 modern floral distributions and subfossil pollen patterns from adjacent regions (e.g., Torres Strait, 175 Atherton Tableland). Extrapolations suggest that similar climatic influences acting on a common flora 176 drove cycles of vicariance of closed forest and expansion of drier woodland in the LB region during 177 the Tertiary (Kershaw, 1985; Rowe 2007). Current vegetation and climate data (Figure 2c‐e) illustrate 178 clearly that the Laura Lowlands are drier and warmer on average during the driest and warmest 179 quarters than the adjacent uplands and currently do not support closed forest refugia. This presents 180 potentially both a climatic and altitudinal barrier to rainforest species on either side of the Laura 181 Lowlands. 182 183 The definition, geographical location and name of the apparent biogeographical barrier for closed 184 forest taxa that occurs in the LB region have been inconsistent in the scientific literature. Three 185 synonymous names have been used to refer to this barrier in the past: the Laura Basin, the 186 Normanby Gap/Barrier and the Torresian Barrier. The Torresian Barrier originally was proposed in a 187 popular book on nocturnal birds by Schodde & Mason (1980) to delineate an apparent disjunction in 188 species distributions at the southern end of Cape York, running in a roughly east‐west direction from 189 Cooktown to Normanton. In reviewing and re‐naming biogeographical barriers to the avifauna Ford 190 (1986) remarked that the name ‘Torresian Barrier’ was inappropriate “for the barrier is neither a 191 major divisor nor a delimitor of Torresian avifaunal elements but effectively marks the southern 192 boundary of Irian elements and the extreme northern limit of Bassian elements” (p. 88). Instead, Ford 193 (1986) preferred to incorporate the area surrounding the “relatively dry Normanby lowlands”, and so 194 renamed this barrier the Normanby Barrier as an alternative name to the Torresian Barrier. The 195 “Normanby lowlands” to which Ford refers must relate instead to what is now known as the Laura 196 Lowlands bioregion, which itself incorporates the Normanby River sub‐basin (Figure 2a) (Wellman, 197 1995; QLD DEHP, 2014). Nevertheless, Ford’s (1986) map (Figure 1, pg 88) showed his Normanby 198 Barrier as a single line running northeast‐southwest from midway between Cooktown and Cape 199 Melville (Figure 2a), despite the “Normanby lowlands” – even if referring only to the Normanby River 200 catchment – in reality occurring to the west of Cape Melville and not reaching the eastern coastline. 201 Ambiguities in nomenclature and location (e.g., Macqueen et al., 2012, Eldridge et al., 2011), 202 including even placement of the barrier south of Cooktown (Edwards & Melville, 2010; Figure 2a), 203 represent a challenge for comparability across the literature. 204 205 Although there is a substantial gap in sampling in this region, despite it being an area of high 206 phylogenetic diversity (Rosauer et al., 2009; Hoskin & Couper, 2013), it appears that Cape Melville is 207 comprised of both northern and southern elements. For example, the northern distributional limit of 208 a species of closed forest orchid occurs at Cape Melville (Burke et al., 2013). Past connection with 209 southern closed forests is also suggested for some geckos (Hoskin & Couper, 2013), skinks (Hoskin, 210 2013a) and frogs (Hoskin, 2013b), in which endemic Cape Melville species were placed in genera 211 otherwise distributed in the Wet Tropics. In contrast, the southern limit of the native rodent 212 Melomys capensis occurs here and the northern limit of its sister taxon M. cervinipes is further south 213 at Cape Flattery (Bryant & Fuller, 2014), with some evidence of genetic structure between Cape 214 Melville and McIlwraith Range M. capensis populations (Bryant, 2013). Cape Melville may thus 215 represent a narrow zone of overlap between northern and southern biotas that does not extend to 216 the closed forest habitats immediately northwest (Iron/McIlwraith Ranges) or south (Daintree), and 217 that the drier zones that separate these regions have limited further dispersal of colonising taxa. 218 Perhaps Cape Melville was not a significant long‐term refuge for closed forest taxa and has been 219 recolonised more recently from north and south, with cycles of isolation on the Cape driving 220 divergence and endemism. On this basis, we recommend future studies consider cautiously the 221 entire Laura Basin region as representing this putative biogeographical barrier, encompassing Cape 222 Melville within the barrier’s boundaries and extending to the east coast south of the Cape, rather 223 than as a narrow barrier invoked for one or the other side of Cape Melville. This is particularly 224 important for forming initial hypotheses in studies that include Cape Melville and allow better 225 characterisation of the biogeographical history of the area. 226 227 Notwithstanding, the LB is considered to have been important in both limiting the modern 228 distributions of southward‐colonising mesic forest species and driving divergence among 229 populations of species that span the barrier (Lavarack & Godwin, 1987). Southward‐colonisers 230 include palm cockatoos (Murphy, Double & Legge, 2007), cuscuses (Winter & Leung, 2008), rodents 231 (Bryant et al., 2011), and pythons (Wilson & Heinsohn, 2007). Most such species apparently 232 colonised Australia from New Guinea via the Torres Strait landbridge during Plio‐Pleistocene glacial 233 cycles before being isolated on Cape York by rising sea levels (see Bryant & Fuller 2014). Possibly, 234 however, their southward colonisation was limited not by the LB, but by other ecological/climatic 235 factors such as temperature or food resources. Several mammal and bird species show either 236 distributions restricted to either side of the LB (Keast, 1961; Ford, 1986) or significant population‐ 237 level genetic/morphological structuring across the barrier (Table 1), and most studies invoke 238 Pleistocene climatic change and associated vicariance of closed forests to explain resolved patterns. 239 In contrast, the orchid D. tetragonum (Burke et al., 2013) and the Central‐Eastern lineage of 240 Leptomyrmex spider ants (Lucky et al., 2011), which have their northern limits at the McIlwraith 241 Range, do not exhibit genetic structuring across the LB implying some degree of dispersal across the 242 barrier. Furthermore, molecular divergence time estimates across the LB extend from the late 243 Pleistocene into the early Miocene (Figure 3), suggesting this barrier may be older than assumed 244 previously. 245 246 Black Mountain Corridor 247 248 Perhaps the best‐characterised biogeographical barrier in eastern Australia is the Black Mountain 249 Corridor (BMC, sometimes referred to as the Black Mountain Barrier – Figure 4a). According to 250 palynological evidence, the BMC historically was an expanse of dry sclerophyll forest that bisected 251 rainforest assemblages in the Daintree and Atherton Tablelands, reaching its maximum extent 252 during the last glacial maximum (LGM) around 38kya (Kershaw, 1985; Bell et al., 1987; Kershaw, 253 1994). Palaeovegetation reconstructions universally imply the region to have been devoid of closed 254 forest or woodland during the LGM, with re‐expansion of rainforest assemblages occurring only in 255 the last 9kya (Kershaw, 1985; Nix & Switzer, 1991; Hopkins et al., 1993). Moreover, pollen evidence 256 for cyclical rainforest contractions in the Wet Tropics is known from as early as 150kya (Kershaw, 257 1994), although little palaeovegetation data precedes this period. The current topography of the 258 region is highly correlated with underlying geology, with upland areas composed mostly of granite 259 substrate and lowlands of metamorphic strata that erode more easily (Nott et al. 2001). Today the 260 BMC is lower in elevation and generally warmer and drier than neighbouring upland rainforest 261 isolates, with a narrow coastal strip of particularly low elevation that supports lowland rainforest 262 possibly because of higher rainfall than the adjacent upland (Figure 4b‐e). 263 264 Phylogeographic studies of a plethora of closed forest taxa generally support a hypothesis of 265 vicariance among northern and southern assemblages, with most studied taxa exhibiting significant 266 genetic breaks across the dry BMC; including, lizards, birds, mammals, invertebrates and plants 267 (Table 1). Several taxa show evidence of re‐expansion and secondary contact of divergent lineages, 268 although dispersal across the BMC seems to have been unidirectional, with sympatry of lineages 269 only on the southern side of the barrier (e.g., Pope, Estoup & Moritz, 2000; Moritz et al., 2009; 270 Krosch et al., 2009). Some other groups that are divergent across the BMC also show significant 271 structure among populations within refugia, particularly to the south (e.g., Atherton Tableland), 272 suggesting that these represent areas of long‐term persistence of closed forest (Moussalli et al., 273 2009, Mellick, Wilson & Rossetto, 2014). Furthermore, patterns in an agamid lizard that inhabits 274 adjacent dry woodland exhibit the expected inverse effect of expansion of drier habitat in this region 275 with evidence for stepping‐stone historical population expansion (Edwards & Melville, 2010). In 276 contrast, some birds (Joseph et al. 1993; Joseph & Moritz, 1994; Joseph, Moritz & Hugall, 1995; 277 Nicholls & Austin, 2005) and mammals (MacQueen et al. 2012; Bryant & Fuller 2014) show no 278 apparent influence of the BMC on population structure (even when they were significantly 279 structured by other nearby barriers), implying that responses to this barrier were species‐specific 280 and over‐generalisation of patterns and effects may be inappropriate. 281 282 Despite broad concordance of significant genetic structure across the BMC across many studies and 283 many taxa, to date relatively few studies have estimated divergence times across the BMC. Of those 284 that have, only divergences within Elaeocarpus species (0.04‐0.18Mya – Mellick et al., 2014) and 285 Carlia rubrigularis (Dolman & Moritz 2006) fall during the late Pleistocene, coinciding directly with 286 fossil pollen evidence for closed forest vicariance. Most estimates, however, range from late 287 Pleistocene to mid‐Miocene, with a distinct outlier estimated for earthworms in the Eocene‐ 288 Cretaceous (31‐84Mya – Moreau et al., 2015) (Figure 3). Within particular groups of taxa, divergence 289 estimates do not overlap among invertebrate and frog taxa, even for species within the frog genus 290 Litoria, whereas divergences for Lampropholis and Diporiphora lizards did overlap. Clearly, the BMC 291 was temporally variable throughout its history (Byrne et al., 2011): direct palynological evidence 292 points to a complex recent history of repeated cycles of contraction of closed forest refugia and 293 expansion of sclerophyll forest. Variation in genetic divergence times among taxa and evidence of 294 secondary contact supports the notion of multiple vicariance events that drove divergence among 295 populations at different points in time. In some taxa, divergence occurred early and proceeding dry 296 periods reinforced vicariance and genetic divergence, while any subsequent recontact of lineages 297 was insufficient to re‐establish homogeneity (Hoskin et al., 2005). In other taxa, however, divergence 298 was much more recent and genetic distances between lineages are more shallow. Future studies 299 must include rigorous dating analyses because divergence in this region cannot be assumed to have 300 occurred synchronously in all taxa. 301 302 Burdekin Gap 303 304 Arguably the largest dryland biogeographical barrier on Australia’s east coast is the Burdekin Gap 305 (BG). The BG coincides with the northern extent of the ‘Brigalow Belt North’ bioregion 306 (Commonwealth of Australia, 2012), a vast dry corridor that stretches from the southern boundary 307 of both the Wet Tropics bioregion and what is considered the Australian Monsoonal Tropics (Catullo 308 et al., 2014), to Bowen in the south (Figure 5a). The region encompasses the dry lowland savannah 309 of the Burdekin River catchment from which the barrier takes its name (Figure 5b,c). The BG 310 separates the remnant upland closed forests of the Paluma Range and Mount Elliot to the north 311 from those in the Clarke Range and Conway Peninsula to the south, with the Clarke Range in 312 particular considered an area of long‐term mesic forest persistence (Stuart‐Fox et al., 2001). The 313 upland refugia are characterised by cooler and wetter climates than the lowlands (Figure 5d,e) and 314 there is currently very little closed forest habitat within the putative bounds of the BG, providing 315 limited opportunities for taxa to disperse across the barrier using smaller patches of suitable habitat 316 as stepping stones. The BG was recognised first and named as such from discontinuities in 317 distributions of bird subspecies and races (Keast, 1961; Galbraith, 1969). It was only after these 318 initial assertions that the region may have constituted an historical barrier that fossil pollen data 319 from adjacent areas was brought to light. This palynological evidence demonstrated that dramatic 320 changes to closed forest distributions had occurred in the late Pleistocene (e.g., Kershaw, 1975), 321 events that were connected subsequently to bird distribution data (Ford 1978, 1979, 1987a,b). 322 However, palaeovegetation composition throughout the BG lowlands is poorly characterised and 323 floral evidence for the past duration and extent of separation/connection of closed forest refugia on 324 either side remains limited. 325 326 Nevertheless, the BG apparently has influenced distribution patterns and driven divergence among 327 populations of many taxa; including, lizards, frogs, birds and mammals (Table 1). Inferred divergence 328 ages from molecular studies show strong concordance, with the majority ranging from the very late 329 Miocene to the late Pleistocene, and an outlier for Uperoleia frogs estimated for mid‐Miocene to 330 Oligocene (Figure 3). All mammal divergences overlap in the mid‐Pleistocene, whereas lizard 331 divergences vary widely in age with minimal overlap (Figure 3). Some species inferred to have 332 diverged across the BG show patterns of subsequent recontact, resulting apparently from uni‐ 333 directional dispersal northward into Mount Elliot and the southern portion of the Paluma Range 334 (Ferraro 2012; Bryant & Fuller 2014). In contrast, no differentiation across the BG has been reported 335 in some taxa, despite the relative importance of other potential barriers (Keleman & Moritz, 1999; 336 Schiffer et al., 2007; Macqueen et al., 2012). Furthermore, although some bird species do show 337 population‐level diversification across the BG (Joseph et al., 1993), it appears that the barrier has 338 been more important in limiting distributions and driving divergence at the subspecies level (Keast, 339 1961; Nicholls & Austin, 2005). It is unclear in how many cases this pattern has resulted from deep 340 divergence between ancestral populations that were isolated by the BG versus latitudinal limits in 341 distribution at the BG driven by thermal or ecological tolerances. This pattern of species distribution 342 limits coinciding with the BG has been observed also in snails (Hugall & Stanisic, 2011), mayflies 343 (Christidis & Dean, 2008; Webb & Suter, 2010) and odonates (Watson & Theischinger, 1984, cited in 344 Webb & Suter 2010). Moreover, deep genetic divergences in some freshwater taxa apparently 345 associated with the BG may in fact have originated with altered drainage boundaries due to Pliocene 346 erosion (Todd et al., 2013; Todd, Blair & Jerry, 2014). Clearly, the BG and adjacent regions have a 347 complex geological, climatic and vegetation history, and significant work will be needed to untangle 348 the relative roles of several biogeographical events on the evolution of the region and its biota. 349 350 Saint Lawrence Gap 351 352 The Saint Lawrence Gap (StLG) is a smaller stretch of low‐lying open woodland and savannah 353 between Mackay and Rockhampton in mid‐eastern Queensland approximately 350 km south of the 354 BG (Figure 5a‐c). The region corresponds to the southeastern portion of the ‘Brigalow Belt North’ 355 bioregion (Commonwealth of Australia, 2012). Although originally referred to as the Broad Sound 356 Barrier, the name Saint Lawrence Gap has been used much more extensively since. We use the 357 latter, both for consistency with current research and because Broad Sound refers to a bay within 358 the putative boundary of the StLG, and an island group and maritime channel offshore, whereas the 359 coastal township of Saint Lawrence is located within the bounds of the barrier. As with the LB and 360 BG this putative biogeographical barrier was recognised first through analysis of avian hybrid zones, 361 in which many avian subspecies appeared to show a disjunction in this region (Keast, 1961; Ford, 362 1986). Typically, these patterns are inferred as the result of historical glacial cycles that drove 363 vicariance of previously continuous populations across the barrier, with little interchange since. 364 Certainly, extant habitat in the region is dominated by dry, open sclerophyll woodland and 365 savannah, with no significant stands of closed forest within the putative boundary of the barrier 366 (Figure 5c); however, palaeovegetation data for the StLG is lacking. The StLG is a noted floristic 367 transition zone, marking the southern and northern boundaries of the tropical and sub‐tropical 368 floras, respectively, and the supposed long‐term geological stability of the region is thought to have 369 allowed mesic communities on either side to evolve independently (Webb & Tracey, 1981). Local 370 climate has been relatively well characterised (Webb & Tracey, 1981), and the StLG appears 371 generally warmer and drier than adjacent uplands (Figure 5d,e). 372 373 Several phylogeographical studies invoke the StLG as an important historical barrier to dispersal 374 among isolated populations, with deep genetic divergences observed across the barrier in several 375 species of skinks, birds, pademelons, insects and orchids (Table 1). In contrast, the evolutionary 376 histories of some other taxa appear less influenced by the StLG (e.g., Toon et al., 2007), despite 377 some exhibiting significant divergence across other eastern Australian biogeographical barriers (e.g., 378 Joseph & Moritz, 1994; Hugall, Stanisic & Moritz, 2003; Bryant & Fuller, 2014). Where structure is 379 observed across the StLG, divergences are generally estimated to be of late Miocene‐late 380 Pleistocene age (Figure 3); the exception being Rix & Harvey (2012), who estimated divergences 381 among assassin spider clades to be of Eocene age (31‐51Mya, Table 1). This is a marked outlier 382 relative to other published estimates for the StLG (Figure 3). This temporal variation in divergence 383 times among taxa implies that multiple biogeographical events have driven several phases of 384 divergence and diversification across the barrier. The StLG is less studied relative to other east coast 385 biogeographical barriers and our full understanding of biotic evolution in this region will benefit 386 from further research into phylogeographical patterns and vegetation history. 387 388 Brisbane Valley Barrier 389 390 Southeast Queensland and northeast New South Wales (NSW) is a complex region for interpreting 391 patterns in closed versus open forest taxa. Topographically, the Brisbane Valley (Figure 6a,b) 392 comprises lowlands encircled by mountain ranges to the near west and south and the Pacific Ocean 393 to the east, and the closed forests that hug the range tops harbour high diversity and endemism 394 (Crisp et al., 2001; Figure 6). The southwestern mountains (Main Range) form part of the GDR, and 395 extending eastward from Main Range to the coast is an east‐west spur of the GDR known as the 396 Border Range, along with the northern portion of the Tweed caldera referred to as the McPherson 397 Range (often misspelt as MacPherson) (Willmott, 2004). To the north, two parallel mountain ranges 398 run south‐north (the GDR to the west and the D’Aguilar Range to the east) that coalesce at the 399 Conondale Range. Climatically, the Brisbane Valley is characterised by being warmer and drier than 400 the adjacent upland regions, and closed forests are limited to high elevation patches on the range 401 tops. These upland refugia are supported as having been highly stable throughout the Pleistocene 402 period compared with intervening lowlands (Weber et al., 2014). Although nomenclature varies 403 among the few studies that have recognised the Brisbane Valley as the barrier to closed forest taxa 404 (e.g., Brisbane River valley – McGuigan et al., 1998; Brisbane Valley Gap – Rix & Harvey, 2012), we 405 prefer to use its most recent incarnation as the Brisbane Valley Barrier (BVB) (Bryant & Fuller, 2014). 406 407 In the broader phylogeographical literature for Australia’s eastern forests, the Main, McPherson and 408 Border Ranges (MMBR) are inferred to have driven the disjunction of open and dry‐forest adapted 409 taxa that inhabit the lowlands either side (e.g., James & Moritz, 2000; Chapple, Chapple & 410 Thompson, 2011). This is intuitive, because these mountains harbour some of the oldest and most 411 stable closed forests outside of the Wet Tropics of north Queensland (Weber et al., 2013; Rosauer et 412 al., 2015) and this, along with their physical topography, has surely acted as a significant ecological 413 barrier to many dry forest taxa in the past. Somewhat confusingly, however, some studies have also 414 considered the MMBR as historical barriers to dispersal of closed forest species, even when extant 415 populations from these ranges are sampled (e.g., Chapple et al., 2011a). Several phylogeographical 416 analyses suggest that closed forest MMBR populations differ significantly from those that inhabit the 417 D’Aguilar and Conondale Ranges further north; including, reptiles, frogs, rodents and arthropods 418 (Table 1). Moreover, when placed in a broader regional context, D’Aguilar and Conondale Range 419 populations are generally more closely related to other populations in mid‐east Queensland, 420 whereas MMBR populations have closer affinities to populations further south. Divergence time 421 estimates across the BVB generally cluster around the Plio‐Pleistocene, except for the much older 422 divergences inferred for Leptomyrmex spider ants (Lucky 2011) and Austrarchaea assassin spiders 423 (Rix & Harvey, 2012). There is clear concordance in age estimates among mammal species – which 424 overlap in the late Pleistocene – whereas estimates for only some lizards overlap (Figure 3). Some 425 taxa appear to have experienced secondary contact across the barrier following initial divergence 426 (e.g., Chapple et al., 2011a), sometimes involving uni‐directional movement southward (e.g., 427 Smissen et al., 2013) or introgression among lineages (e.g., Bryant & Fuller, 2014). Furthermore, 428 several freshwater taxa exhibit significant genetic structure among the Brisbane and Mary River 429 drainages north of the BVB associated with a single divergence event during the Pleistocene, 430 suggesting potential ‘refugia within refugia’ (Hodges, Donnellan & Georges, 2014; Page & Hughes 431 2014). Another apparent zone of transition between biotas in this region occurs around the Clarence 432 River Valley, at the very southern end of the MMO, known as the Clarence River Corridor (CRC, 433 Figure 6) (Mellick, Lowe & Rossetto, 2011; Mellick et al., 2012). The CRC appears to have been 434 important in the diversification of several closed forest geckos (Couper et al., 2008; Colgan, O’Meally 435 & Sadlier, 2009), fish (Rourke & Gilligan, 2010), plants (Mellick et al., 2011, 2012; Heslewood et al., 436 2014; Van Der Merwe et al., 2014), and mammals (Rowe et al., 2012; Frankham, Handasyde & 437 Eldridge, 2012). Likewise, some taxa appear to be restricted by the intersection of open woodland 438 around the Glasshouse Mountains region north of Brisbane (Crisp, Linder & Weston, 1995). 439 440 Taken together, upland regions that have sustained significant stable closed forest refugia through 441 time like the MMBR are unlikely to have simultaneously been important biogeographical barriers for 442 closed forest endemic taxa. Instead, evidence is stronger that the dry lowland Brisbane Valley 443 represents a barrier for many closed forest taxa, and the MMBR are important as barriers 444 predominately for dry forest‐adapted taxa or those that are limited to lower elevations. These two 445 barriers combined have contributed to the broader McPherson‐Macleay Overlap zone (MMO, 446 sometimes called the Macleay‐McPherson Overlap), of which southeast Queensland represents the 447 northern portion (Figure 6). This region was recognised originally for the gradual transition from 448 subtropical to temperate floral communities that occurs across southeast Queensland and northeast 449 NSW (Burbidge, 1960). The MMO is defined as extending from 50km north of Brisbane to the Hunter 450 Valley in NSW (Ebach et al., 2013). This latitudinal transition in regional floras is likely to be 451 reinforced by climatic trends. Burbidge (1960) argued that the marked winter dry season typical of 452 regions north of the MMO limited temperate species’ ability to produce winter growth and so 453 compete with subtropical species. In contrast, subtropical taxa appear to face no such restriction to 454 dispersal southward. Interestingly, regions north and south of the MMO receive high mean annual 455 rainfall, whereas lower rainfall occurs in the MMO itself (Weber et al., 2014). It appears, then, that 456 the CRC, BVB and MMBR have each contributed to the north‐south geographical spread of genetic 457 subdivisions and species distributional limits across southeast Queensland and northern NSW, and 458 which combined seems to have created the broader MMO (Burbidge, 1960). Distinguishing between 459 historical barriers and which taxa they are likely to have influenced is important to understand the 460 history of this region better and we urge future workers to consider carefully the barriers most 461 relevant to their target organism. We stress the importance of an appropriate sampling design to 462 test adequately the relative influence of these barriers. Moreover, better characterisation of the 463 affinities of the less well sampled closed forest refugia adjacent to or within the Brisbane Valley (e.g., 464 at Crows Nest and Mount Tamborine) will help to clarify the historical impact of the BVB. 465 466 Hunter Valley Barrier 467 468 The Hunter Valley Barrier (HVB) is located in eastern NSW and comprises extensive lowlands flanking 469 the Hunter River, which flows southwest from montane Barrington Tops before coalescing with the 470 east‐flowing Goulburn River and turning southeast until it reaches the coast at Newcastle (Figure 471 7a,b). We incorporate the Goulburn Valley lowlands into our concept of the broader Hunter Valley 472 hereafter, in keeping with the majority of phylogeographical literature. The Hunter Valley was first 473 proposed as a potential barrier to north‐south dispersal for several Eucalyptus species (Cameron, 474 1935) and has been invoked as delineating the southern limit of an ‘Eastern Biogeographic Region’ 475 based on early analyses of avian species distributions (Ford, 1987a,b; Cracraft, 1991). This hypothesis 476 has been supported by more recent modelling of species endemism, turnover and habitat stability 477 reveal an abrupt break between assemblages north and south of the barrier, corresponding with the 478 boundaries of the Sydney Basin and the Hunter Valley (Di Virgilio, Laffan & Ebach, 2012; Rosauer et 479 al., 2015). Disjunctions across the HVB region have occasionally been ascribed to the Cassilis Gap 480 (e.g., Rix & Harvey, 2012), a lower elevation pass through the GDR at the head of the Goulburn River 481 Valley that is thought to separate the Liverpool Range from the northern Blue Mountains (Sussmilch, 482 1929; Cotton, 1949) (Figure 7a). The Cassilis Gap is geologically more ancient than the proposed 483 Neogene‐Quaternary contraction of closed forests and is thought to have facilitated movement of 484 organisms from one side of the range to the other (Hobbs & Kaveney, 1962; Moore, 1970; Davies, 485 1977; Hodges, Donnellan & Georges, 2015). Nevertheless, the Cassilis Gap and the Hunter Valley 486 lowlands may have acted in concert to impede longitudinal dispersal of closed forest taxa otherwise 487 restricted to the mountaintops of the GDR and we consider them together here. The Hunter Valley is 488 characterised by dry, open woodland and savannah that bisects upland closed forest habitat to the 489 north (e.g., Barrington Tops) and south (e.g., Blue Mountains) and is typically warmer and drier than 490 the adjacent uplands (Sweller & Martin, 2001, Figure 7c‐e). 491 492 The HVB has been implicated in limiting species distributions and driving speciation in a range of 493 closed forest taxa; including, snails, assassin spiders and mayflies (Table 1). Intraspecific patterns 494 suggest similar processes acting at the population level in several skink, plant, butterfly and 495 marsupial species. As with some of the other putative barriers discussed above, the HVB or the 496 broader region surrounding it, has been implicated as also influencing divergence among 497 populations of some open forest‐restricted taxa; including, frogs and lizards (Table 1). It remains 498 unknown how a region of open forest should act as a biogeographical barrier to taxa that otherwise 499 might inhabit it: perhaps other processes have driven diversification of these taxa in this region. 500 Furthermore, there is some variation in the responses of studied closed forest taxa to this putative 501 gap, with numerous mammals, lizards and frogs showing no structure across the HVB (Table 1). In 502 some taxa, disjunctions among clades occur further south of the Hunter Valley, possibly associated 503 with other lowland open forests (see following section), although this pattern may also represent 504 southward re‐expansion of northern populations following initial divergence across the HVB. 505 Estimated divergence times generally are concordant with expectations for vicariance of closed 506 forest driving divergence in isolation, appearing to cluster around the Pliocene, with older 507 divergences estimated for Lampropholis guichenoti in the mid‐Miocene (Chapple et al., 2011b) and 508 late Miocene for Austrarchaea assassin spiders (Rix & Harvey, 2012) (Figure 3). Although many 509 questions remain concerning the impact and variation in the role of the putative HVB, this region has 510 evidently limited movement among populations in many groups. The southern temperate closed 511 forests are considerably understudied relative to the tropical northern regions (Chapple et al. 2011b) 512 and it is likely that continued research into phylogeographical patterns among closed forest taxa 513 from this region will shed further light on the biogeographical processes that have shaped the 514 regional biota. 515 516 Southern Transition Zone 517 518 Southeastern NSW features an extensive north‐south band of lower elevation open woodland and 519 grassland that extends from west of the Illawarra region south nearly to the NSW‐Victorian border 520 (Figure 8a,b). The northern portion of the STZ separates the Australian Alps and the Blue Mountains/ 521 Southern Highlands at the geographical feature known as the Lake George Gate (Taylor 1914, 1959; 522 Karskens 2014), whereas the southern end divides the lowland coastal forests from the higher 523 elevation montane habitats of the GDR. Several historical events have shaped the biogeography of 524 the region: Cretaceous‐Neogene tectonic uplift of the GDR that bisected populations east and west; 525 widespread volcanism and lava flows during orogeny may have fragmented habitats further; 526 subsequent erosion opened the lowland Lake George Gate, isolating mountaintop populations; Mio‐ 527 Pleistocene glaciations drove retraction of closed forest into cool, wet gullies isolated by drier 528 lowlands and higher elevation alpine zones (Wellman & McDougall, 1974). Currently, the region 529 experiences markedly lower rainfall than both the alpine regions to the southwest, the lowland 530 coastal zone to the east, and the Blue Mountains to the northeast (Figure 8e). There is a distinct 531 difference in floral communities, with high diversity and endemism in the Blue Mountains/Sydney 532 Basin region (Crisp et al., 2001) and markedly higher species turnover southward across the STZ (Di 533 Virgilio et al., 2012), although compositional turnover in frog and lizard lineages occurs further north 534 in the Blue Mountains/Sydney Basin region (Rosauer et al., 2015). 535 536 The STZ appears to have been important in driving Miocene‐Pleistocene divergence in a variety of 537 closed forest taxa; including, invertebrates, lizards, birds, frogs, mammals and plants (Table 1). 538 Disjunctions between divergent genetic clades generally are located toward the southern end of the 539 STZ, with lowland coastal populations immediately east appearing divergent to populations on the 540 slopes of the Australian Alps and/or eastern coastal Victoria. Divergence time estimates for lizards 541 largely overlap in the late Miocene to early Pleistocene (Figure 3), but assassin spiders appear to 542 have diverged across the STZ in the Paleocene‐Eocene (Rix & Harvey, 2012). In the latter case, 543 orogeny and volcanism associated with uplift of the GDR probably were most important in driving 544 divergence, whereas divergences in other taxa coincide with Miocene climatic change and forest 545 vicariance. At a finer geographical scale (~100km north‐south) within this zone, many taxa in the 546 Tallaganda region (Figure 8a) exhibit significant latitudinal structuring of Mio‐Pleistocene age 547 associated with microgeographic zones delimited by topography and historical rainfall regimes 548 (Garrick et al., 2004; Hodges, Rowell & Keogh, 2007; Beavis, Sunnucks & Rowell, 2011; Garrick, 549 Rowell & Sunnucks, 2012). It is likely that part of the Tallaganda region acted as a refuge for mesic 550 taxa within the broader STZ; however, some uncertainty exists surrounding exactly what processes 551 might be responsible for maintaining divergence on such a small scale. 552 553 Other regions nearby the STZ that have been implicated as important in structuring genetic diversity 554 within species, albeit with less concordance across taxa, are the Sydney Basin and the Illawarra 555 region (Figure 8a). Both regions lay to the northeast of the STZ: the Sydney Basin representing the 556 lowlands surrounding Sydney, and the Illawarra proximately south of this. The regions are 557 characterised by low‐lying coastal plains bounded to the west by the Blue Mountains/Southern 558 Highlands and the Illawarra Escarpment, a region of uplifted sandstone that forms the southern end 559 of the Southern Highlands. Much of the Sydney Basin has been heavily cleared; however, the 560 Illawarra region is characterised by open forest types that almost entirely separate coastal closed 561 forests (Figure 8c), and it is reasonable to expect that closed forests would have been extirpated 562 from these regions during historical dry periods. Several studies suggest that one or other of these 563 regions may have acted as a barrier to some species of closed forest reptiles (Sumner et al., 2010; 564 Dubey et al., 2012; Pepper et al., 2014), assassin spiders (Rix & Harvey, 2012), and mammals 565 (Frankham, Handasyde & Eldridge, 2015), although sampling designs were often unable to 566 distinguish between the two. Along with probable impacts of Mio‐Pleistocene vicariance of closed 567 forests, the sandstone Escarpment is also considered an ecological barrier for some snakes because 568 it is unsuitable to support thermoregulation (Sumner et al., 2010). Taken together, the STZ and 569 adjacent regions have a diverse biogeographic history and is supported as a topographical barrier for 570 alpine species across the Lake George Gate region, and a climatic/ecological barrier for taxa from the 571 adjacent closed forests. 572 573 East Gippsland Barrier 574 575 The final putative barrier to eastern Australian closed forest taxa treated here is the East Gippsland 576 Basin (EGB) in eastern Victoria (Figure 8). The Gippsland Basin is a region of open‐forested lowlands 577 in southeastern Victoria that is bounded to the north and west by the southern GDR (Australian 578 Alps) with closed forests restricted to the emergent Strzelecki Range in the centre of the Basin. 579 Although the EGB doesn’t appear distinctly warmer than the Strzelecki Range, it is more warm and 580 distinctly drier than both the closed forests of the GDR and the coastal regions to the northwest 581 (Figure 8d,e). Extensive tectonic uplift occurred at the southern GDR from the Eocene‐Oligocene 582 (Holdgate et al., 2008), and the lowlands of the EGB were submerged during periods of higher sea 583 level since the Miocene (Lambeck & Chappell, 2001; Dickinson et al., 2002). Some parts of the Basin 584 were likely also to have been glaciated during the Miocene‐Pleistocene (MacPhail, Colhoun & 585 Fitzsimons, 1995), driving transition from Araucaria and Nothofagus‐dominated forests to more 586 open, dry woodlands (Truswell 1993; Gallagher et al., 2003). The EGB has been implicated in Upper 587 Cretaceous‐Paleocene speciation in assassin spiders (Rix & Harvey, 2012), which is likely related to 588 orogeny and volcanism associated with uplift of the GDR. A number of studies also support Mio‐ 589 Pliocene diversification across the EGB in some lizards, frogs, and plants; however, many taxa do not 590 exhibit structure here (Table 1). The drivers of these Mio‐Pliocene divergences have been difficult to 591 untangle: both closed forest vicariance resulting from aridification of the lowlands, and inundation 592 of the lowlands leaving closed forests isolated on islands, presumably leave similar legacies in the 593 population structure of taxa of the region. This region thus presents some intriguing questions 594 concerning the relative impact of biogeographical events on taxa which can be resolved only via 595 additional phylogeographical assessments. 596 597 Common trends among eastern Australian barriers 598 599 This review points to some clear trends among the major eastern Australian biogeographical barriers 600 and associated biotic transition zones that dissect remnant closed forest habitats along the eastern 601 Australian seaboard. All eight reviewed barriers are lower elevation zones of dry, warm, open 602 woodland/grassland habitat that are unsuitable for closed forest taxa. In almost every case, closed 603 forests adjacent to these barriers occupy cooler and wetter uplands compared with the intervening 604 lowlands. An exception is the Daintree coastal rainforest north of the Black Mountain Corridor, 605 although it too encompasses upland habitats and experiences significant seasonality in monsoonal 606 rainfall. Furthermore, molecular divergence estimates among the barriers fall largely within the last 607 25Mya (Miocene‐present), with particular peaks in simultaneous divergence at multiple barriers 608 around the mid‐late Miocene (~15Mya) and the Pliocene‐Pleistocene (~6‐0.04Mya) (Figure 2). 609 Drawing direct comparisons between studies can be misleading, because different loci, molecular 610 clock rates, calibration points and analytical methods used across studies may give vastly different 611 estimates that may not reflect reality. However, general convergence of genetic divergence time 612 estimates among such a wide range of taxa and concordant with estimates for the onset of 613 vicariance of widespread closed forests is striking. 614 615 Perhaps unsurprisingly, these eight regions represent the majority of significant breaks in the current 616 distribution of closed forests in eastern Australia and modelling analyses of species turnover do not 617 indicate distinct transition zones or biotic breaks occurring at any other locations along the east 618 coast (e.g., Rosauer et al., 2015). Only the areas between the Jardine River headwaters and the Iron 619 Range on Cape York, and between the southern end of the Saint Lawrence Gap and the Conondale 620 Range (associated especially with the Burnett River basin) appear to have similar characteristics to 621 the reviewed barriers, but apparently have not influenced population genetic structure as 622 dramatically. Other low‐elevation zones that separate upland regions ‐ for example, the Sydney 623 Basin, northeastern NSW – do not possess the same vegetation or climatic traits as the major 624 barriers, and appear to have had less influence on population structure across a broad range of taxa. 625 Thus, there was presumably something fundamental about the combination of elevation, climate 626 and resident floral communities at the eight major barriers and the manner in which they were 627 influenced by historical climate change that has created such distinctly concordant biotic patterns. 628 629 Nevertheless, variation exists among taxa in their apparent response to each proposed 630 biogeographical barrier: all eight barriers have been ‘leaky’. Many taxa with distributions that span 631 multiple barriers are not structured across all or any of them (Table 1), suggesting differential 632 influence of some barriers over others for such taxa. A reasonable expectation is that highly 633 dispersive and/or ecologically tolerant taxa may not be influenced by these barriers, but this does 634 not adequately explain why such taxa might freely disperse across one barrier but be highly 635 structured across another. Such scenarios may imply other demographic effects (e.g., local 636 extinction, selective sweeps), as equally plausible in explaining significant genetic structure across 637 barriers as random jump dispersal across unsuitable habitat or lack of fixation of alleles may explain 638 homogeneity. Additionally, the age and geographical extent of a particular biogeographical barrier, 639 the availability of any remnant patches of appropriate habitat within the zone of otherwise 640 unsuitable habitat and the length of time particular taxa have been present in the adjacent region all 641 determine the ‘leakiness’ of a given barrier. Furthermore, for many of the above barriers, re‐ 642 expansion of previously isolated, divergent populations across the barrier appears to have facilitated 643 secondary contact and sympatry of divergent lineages via regular dispersal during periods of closed 644 forest expansion (Phillips et al., 2004; Hoskin et al., 2005; Moritz et al., 2009). Re‐expansion of 645 divergent lineages appears to have been particularly frequent across the BMC and BVB, and is likely 646 also to have given rise to the current species assemblage found at Cape Melville in the midst of the 647 LB. Taken together, observed variation among taxa in their response to a given barrier and their 648 propensity for re‐expansion is likely to have been driven by behavioural, physiological, demographic 649 and autecological factors that have interacted differently with the physical environment itself. 650 651 Prospectus for future research 652 653 In this review we have brought together phylogeographical and paleoecological information for six 654 major postulated dry habitat biogeographical barriers along Australia’s east coast that influenced the 655 evolution, diversification and ecology of numerous closed forest‐restricted taxa. Although this 656 literature base is considerable and our understanding of the actual biogeographical attributes of 657 each barrier is perceived to be robust, there exist abundant opportunities to utilise Australia’s 658 eastern mesic forest archipelago not only as a model system for exploring the processes that drive 659 the evolution of diversity, but also to increase knowledge and awareness of the evolutionary history 660 of Australia’s biota. Some of these questions include resolving the affinities of the Cape Melville 661 biota; better characterising biotic patterns in regions immediately south of the BMC and BG that 662 suggest long‐term areas of persistence and potential ‘refugia within refugia’; resolving patterns of 663 sympatry, introgression and presumed differential dispersal among divergent clades at the northern 664 boundary of the BG (southern Paluma Range); exploring the complex patterns reported around the 665 BVB and broader MMO to shed further light on its complicated vegetation, geological and climatic 666 history, with future studies accounting for this with extensive sampling across the region; and 667 assessing in more detail phylogeographical patterns at the HVB, STZ and EGB. Furthermore, some 668 isolated refugia of high diversity and endemism warrant further investigation; including, Kroombit 669 Tops, the Clarke Range and the Mount Warning Caldera at the Queensland‐NSW border (e.g., 670 Hoskin, Couper & Schneider, 2003; Baker, Mutton & Hines, 2013; Baker et al., 2014; Bryant & Fuller, 671 2014). These regions only now are beginning to reveal the secrets of their evolutionary history. 672 673 Future studies of Australian east coast biogeography should expand the taxonomic coverage of 674 current phlyogeographical knowledge by investigating evolutionary patterns in less well‐known 675 closed forest‐restricted taxonomic groups. For example, the small mammal fauna contains many 676 closed forest endemics with distributions that span biogeographical barriers but have significant 677 gaps that correspond with unsuitable habitat; including, dasyurids, rodents and macropods (Strahan 678 & Van Dyck, 2006). Recent work on Antechinus species has hinted at significant effects of historical 679 habitat fragmentation and population isolation on the diversification and evolution of this genus 680 (Baker, Mutton & Van Dyck, 2012; Baker et al., 2013, 2014). Similarly, little attention has been given 681 to closed forest insects; only a handful of studies exist despite their huge diversity. Perhaps 682 taxonomic uncertainty has limited such work in some insect groups, yet many represent ideal 683 models for investigation (e.g., leaf litter and dead wood taxa). Moreover, critical comparative 684 analyses should be conducted between groups of taxa with different ecologies and physiologies; for 685 example, between solely terrestrial and aquatic taxa, or between taxa with putatively high and low 686 vagility. In particular, implementation of direct comparative statistical approaches which can 687 account for coalescent and demographic variation to analyse multiple taxa simultaneously (e.g., 688 hierarchical approximate Bayesian computation, see Dolman & Joseph, 2012; Page & Hughes, 2014) 689 would provide novel insights into the relative importance of these biogeographical barriers in driving 690 evolution in Australia. 691 692 To complement future phylogeographical studies of eastern closed forest taxa, the field would 693 greatly benefit from additional targeted palynological and paleoecological studies. The story of 694 Australian vegetation change since the Miocene is generally told in a broad continental context (e.g., 695 Martin, 2006) and, although drawing upon a wealth of data, mostly derives from southeastern 696 Australia and the Atherton Tableland. Targeted studies that investigate the specific regional 697 vegetation history at each barrier are largely lacking. Some exceptions include the BMC, for which 698 Pleistocene data from the Atherton Tableland has been of most import (e.g., Kershaw, 1985; Bell et 699 al., 1987; Kershaw, 1994), the StLG where several Tertiary fossil sites around the barrier have been 700 assessed (Hekel, 1972), and the HVB for which data from nearby Barrington Tops informs of late 701 Pleistocene vegetation change (Sweller & Martin, 2001). Excluding Hekel’s (1972) macrofossil data, 702 even these studies do not assess sites within the actual contemporary boundaries of the barriers. 703 Thus, the lack of fine resolution data concerning past changes in closed forest distribution and 704 composition at the sites of the major barriers remains a key knowledge gap for ongoing future 705 phylogeographical research. 706 707 Finally, a critical issue for studies of biogeographical barriers worldwide is to ensure the initial 708 experimental design is adequate to test the impact and timing of a given barrier. This includes 709 implementation of adequate sampling both spatially and with sufficient sample size per population 710 to test the influence of the target barrier most rigorously. Significant sampling gaps can lead to 711 ambiguity concerning the geographical location of any observed genetic break that may be 712 associated with a putative barrier and can mask more subtle isolation by distance patterns and 713 latitudinal clines, because intermediate populations are unsampled. Most importantly, zones of 714 secondary contact between divergent lineages may be overlooked without dense sampling of 715 populations adjacent to a barrier edge (e.g., Chapple et al., 2011a; Smissen et al., 2013; Bryant & 716 Fuller, 2014). Furthermore, molecular studies should estimate divergence times for clades separated 717 by barriers using appropriate calibrations. Many early molecular studies lacked rigorous divergence 718 time estimation and thus patterns of population structure were discussed only in light of available 719 pollen evidence for the late Pleistocene. Recent evidence implies more ancient divergences and it 720 remains crucial that future studies test these hypotheses. Similarly, statistical approaches based on 721 ecological niche models allow hypotheses about the impact of specific barriers to be tested directly 722 and which could be incorporated in the future (Glor & Warren, 2011). 723 724 In conclusion, the archipelagic nature of eastern Australia’s closed forest habitats provides an ideal 725 natural laboratory to explore the impacts of habitat fragmentation on terrestrial populations. We 726 implore the research community to retain interest in this enigmatic system as continuing to reveal 727 the history of life in the eastern forests adds to the patchwork of knowledge about the evolution, 728 origins and diversity of Australia’s biota. Understanding the historical impacts of climate change 729 informs predictions of future expected responses of taxa to human‐induced climate change, 730 accepted as an imminent risk to closed forests, especially in the tropics (Williams, Bolitho & Fox, 731 2003; Williams & Hilbert, 2006), and allows development of effective future management and 732 conservation strategies. 733 734 Acknowledgements 735 736 The authors thank Peter Cranston, Andrew Baker, Susan Fuller, David Hurwood, Peter Mather and 737 Thomas Mutton for many fruitful discussions concerning Australian east coast biogeography. 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Shaded cells denote taxa that are structured across the corresponding barrier, with divergence times given if they were 1245 available or coded by Y if not. Unshaded cells are coded with an X if no influence of the corresponding barrier was inferred, or ‐ if the sampling design did 1246 not encompass the barrier or was insufficient. Barrier & divergence time estimates Species & reference LB BMC BG StLG BVB HVB STZ EGB Mammals Thylogale stigmatica (Eldridge et al., 2011; MacQueen et al., 2012) Y X X Y X ‐ ‐ ‐ Uromys caudimaculatus & Rattus leucopus (Baverstock, Watts & Hogarth, Y ‐ ‐ ‐ ‐ ‐ ‐ ‐ 1977) Melomys capensis & M. cervinipes (Bryant & Fuller, 2014) 0.3‐1.3 ‐ ‐ ‐ ‐ ‐ ‐ ‐ Bettongia tropica (Pope, Estoup & Moritz, 2000) ‐ Y ‐ ‐ ‐ ‐ ‐ ‐ Petaurus australis (Brown et al., 2006; Ferraro, 2012) ‐ ‐ 0.6‐1.4 X X X ‐ ‐ Melomys cervinipes (Bryant et al., 2011; Bryant & Fuller, 2014) ‐ X 0.5‐1.6 X 0.2‐0.8 ‐ ‐ ‐ Petrogale penicillata (Eldridge et al., 2001; Hazlitt et al., 2014) ‐ ‐ ‐ ‐ X Y Y X Phascolarctos cinerus (Houlden et al., 1999) ‐ ‐ ‐ ‐ Y X X Y Dasyurus maculates (Firestone et al., 1999) ‐ ‐ ‐ ‐ ‐ X X ‐ Petrogale penicillata (Potter et al., 2012) ‐ ‐ 0.05‐2.9 0.1‐4.4 0.05‐2.8 X X X Potorous tridactylus (Frankham et al., 2015) ‐ ‐ ‐ ‐ ‐ ‐ Y Y

Birds Melithreptus albogularis (Toon, Hughes & Joseph, 2010) 1.8‐20 X X X X X ‐ ‐ Sericornis magnirostris, Psophodes olivaceus (Joseph et al., 1993) ‐ ‐ Y Y X ‐ ‐ ‐ Ptilonorhynchus violaceus (Nicholls & Austin, 2005) ‐ X Y Y X X X X Gymnorhina tibicen (Toon et al., 2007) ‐ ‐ X X X X X X Poecilodryas albispecularis, Orthonyx spaldingii, Sericornis citreogularis ‐ Y ‐ ‐ ‐ ‐ ‐ ‐ (Joseph et al., 1995; Joseph & Moritz, 1994)

Reptiles Saltuarius cornutus, Carphodactylus laevis, Gnypetoscincus queenslandiae, (Moritz, Joseph & Adams, 1993; Joseph et al., 1995; ‐ Y ‐ ‐ ‐ ‐ ‐ ‐ Cunningham & Moritz, 1998; Schneider, Cunningham & Moritz, 1998; Schneider & Moritz, 1999) Carlia rubrigularis (Schneider et al., 1999; Phillips et al., 2004; Dolman & ‐ 0.8 ‐ ‐ ‐ ‐ ‐ ‐ Moritz, 2006) Lampropholis robertsi, L. coggeri (Bell et al., 2010) ‐ 1.8‐11.7 ‐ ‐ ‐ ‐ ‐ ‐ Carlia rhomboidalis (Dolman & Moritz, 2006) ‐ ‐ 0.3 ‐ ‐ ‐ ‐ ‐ Saproscincus czechurai, S. basiliscus, S. tetradactyla (Moussalli et al., ‐ ‐ Y ‐ ‐ ‐ ‐ ‐ 2009) Diporiphora australis (Edwards & Melville, 2010) ‐ 4.7‐9.2 3.5‐6.9 X 1.3‐3.2 ‐ ‐ ‐ Saproscincus sp. (Moussalli, Hugall & Moritz, 2005) ‐ X Y Y X Y ‐ ‐ Lampropholis delicata (Chapple et al., 2011a) ‐ X 3‐4 3‐4 4‐5 4‐5 2.4‐3.2 ‐ Varanus varius (Smissen et al., 2013) ‐ X 0.06‐1.7 X 1.6‐3.3 X ‐ ‐ Eulamprus sp. (O'Conner & Moritz, 2003) ‐ ‐ X Y X X ‐ ‐ Hoplocephalus stephensi (Keogh et al., 2003) ‐ ‐ ‐ ‐ Y ‐ ‐ ‐ Saproscincus rosei (in Rosauer et al., 2015) ‐ ‐ ‐ ‐ X ‐ ‐ ‐ Lampropholis guichenoti (Chapple et al., 2011b) ‐ ‐ ‐ ‐ 2.2‐5.8 8‐12 X 3.6‐5.8 Bassiana platynota (Dubey & Shine, 2010) ‐ ‐ ‐ ‐ ‐ 2.3‐5.9 1.4‐19.3 ‐ Bassiana duperreyi (Dubey & Shine, 2010) ‐ ‐ ‐ ‐ ‐ ‐ 2.3‐6.6 X Amphibolurus muricatus (Pepper et al., 2014) ‐ ‐ ‐ ‐ ‐ Y X Y Liopholis whitii (as Egernia whitii ‐ Chapple & Keogh, 2004; Chapple et al., ‐ ‐ ‐ ‐ X X X 2.8‐9.6 2005) Rankinia diemensis (Ng et al., 2014) ‐ ‐ ‐ ‐ ‐ ‐ Y ‐ Delma impar (Maldonado et al., 2012) ‐ ‐ ‐ ‐ ‐ ‐ Y ‐

Frogs Litoria genimaculata, L. rheocola, L. nannotis (Schneider et al., 1998) ‐ Y ‐ ‐ ‐ ‐ ‐ ‐ Litoria serrata (Bell et al., 2012) ‐ 7‐12 ‐ ‐ ‐ ‐ ‐ ‐ Litoria jungguy, L. rheocola, L. dayi, L. nannotis (Bell et al., 2012) ‐ 2‐5 ‐ ‐ ‐ ‐ ‐ ‐ Cophixalus sp. (Hoskin et al., 2011) ‐ Y ‐ ‐ ‐ ‐ ‐ ‐ Mixophyes sp. (Oza et al., 2012) ‐ Y ‐ ‐ ‐ ‐ ‐ ‐ Limnodynastes tasmaniensis, L. peronii (Schauble & Moritz, 2001) ‐ ‐ Y X X Y Y X Litoria fallax (James & Moritz, 2000) ‐ ‐ Y X Y X ‐ ‐ Litoria pearsoniana (McGuigan et al., 1998) ‐ ‐ ‐ ‐ Y ‐ ‐ ‐ Litoria citropa sp. grp. (Donnellan et al., 1999) ‐ ‐ ‐ ‐ X Y Y X Litoria aurea (Burns et al., 2004, 2007) ‐ ‐ ‐ ‐ ‐ X Y X Crinia signifera (Symula et al., 2008) ‐ ‐ ‐ ‐ ‐ X 4.5‐11.9 6.8‐11.9 Uperoleia sp. (Catullo & Keogh, 2014; Catullo et al., 2014) ‐ ‐ 9.8‐24.3 ‐ ‐ ‐ ‐ ‐

Fish Pseudomugil signifer (McGlashan & Hughes, 2002; Wong, Keogh & ‐ X Y X X ‐ ‐ ‐ McGlashan, 2004; Kelly et al., 2013)

Invertebrates Temnoplectron sp. (Bell et al., 2004, 2007) ‐ 1.7‐2.7 ‐ ‐ ‐ ‐ ‐ ‐ Echinocladius martini (Krosch et al., 2009; Krosch, 2011) ‐ 6‐7 ‐ ‐ ‐ ‐ ‐ ‐ Batrachomyia sp. (Hoskin & McCallum, 2007) ‐ X ‐ ‐ ‐ ‐ ‐ ‐ Austrophleboides sp. (Christidis & Dean, 2008) ‐ X Y X X Y X X Austropurcellia sp. (Popkin‐Hall & Boyer, 2014) ‐ ‐ Y ‐ ‐ ‐ ‐ ‐ Arachnocampa sp. (Baker et al., 2008) ‐ X 6.2 6.2 4 5.5 3.2 3.5 Leptomyrmex sp. (Lucky, 2011) X X X X 8‐22 X X X Bungona narilla (McLean, Schmidt & Hughes, 2008) ‐ ‐ ‐ ‐ Y ‐ ‐ ‐ Jalmenus evagoras (Eastwood et al., 2006) ‐ ‐ ‐ ‐ ‐ Y X X Terrisswalkerius sp. (Moreau et al., 2015) ‐ 31‐84 ‐ ‐ ‐ ‐ ‐ ‐ Planipapillus sp. (Rockman, Rowell & Tait, 2001) ‐ ‐ ‐ ‐ ‐ ‐ X Y Austrarchaea sp. (Rix & Harvey, 2012) ‐ X X 34‐51 18‐27 16‐23 46‐69 ‐ Euastacus sp. (Shull et al., 2005; Ponniah & Hughes, 2004a,b) ‐ Y X X Y X Y X Gnarosophia bellendenkerensis (Hugall et al., 2002) ‐ Y ‐ ‐ ‐ ‐ ‐ ‐

Plants Dendrobium speciosum (Burke et al., 2013) ‐ 2‐7 X 0.6‐4 X X ‐ ‐ Dendrobium tetragonum (Burke et al., 2013) X X X 2‐6 X X ‐ ‐ Elaeocarpus foveolatus, E. largiflorens & E. carolinae (Rossetto et al., ‐ 0.04‐0.18 ‐ ‐ ‐ ‐ ‐ ‐ 2007, 2009; Mellick et al., 2014) Elaeocarpus angustifolius, E. bancrofti, E. sericopetalus, E. elliffi, E. ‐ X ‐ ‐ ‐ ‐ ‐ ‐ johnsoni, E. ferruginiflorus (Rossetto et al., 2007, 2009) Eucalyptus globulus gp. (Jones et al., 2013) ‐ ‐ ‐ ‐ ‐ ‐ Y X Eucalyptus grandis (Jones et al., 2006) ‐ ‐ ‐ ‐ X ‐ ‐ ‐ Ceratopetalum apetalum (Heslewood et al., 2014) ‐ ‐ ‐ ‐ ‐ Y ‐ ‐ Lomatia sp. (Milner et al., 2012) ‐ ‐ ‐ ‐ ‐ Y ‐ Y Telopea sp. (Rossetto et al., 2012) ‐ ‐ ‐ ‐ ‐ ‐ Y Y 1247 1248 Figure Legends 1249 1250 Figure 1. Relative geographical locations of the eight major eastern Australian dryland 1251 biogeographical barriers discussed here. Putative barriers locations are denoted by the dashed 1252 shapes and bold type names, phytogeographical subregions of Ebach et al. (2013) are enclosed in 1253 the solid lines and names are in italics. 1254 Figure 2. Proposed geographical location of the Laura Basin (LB). Inset a) satellite imagery of the LBB: 1255 extents of the Laura Lowlands bioregion and Normanby River catchment are shown within the 1256 bounds of the small and large dashed lines, respectively. Solid lines indicate some of the previous 1257 published locations of the LB ‐ 1: MacQueen et al. (2012), 2: Ford (1986), Eldridge et al. (2011); 3: 1258 Edwards & Melville (2010). b) elevation (masl); c) International Geosphere‐Biosphere Programme 1259 (IGBP) land cover; d) temperature (warmest quarter mean); e) precipitation (driest quarter mean). In 1260 all figures, locations referred to in the text are indicated by arrows and filled circles with italicised 1261 labels indicate major townships. Satellite images were produced using Google Earth; environmental 1262 layers were produced using the Atlas of Living Australia’s Mapping and Analysis web portal 1263 (http://spatial.ala.org.au/#). 1264 Figure 3. Molecular divergence time estimates from studies assessing phylogeographic patterns at 1265 the eight major eastern Australian barriers, from Table 1. The barrier each divergence relates to is 1266 given in the Y axis labels, using initials as per the text. X’s denote age estimates that lacked 1267 confidence intervals. Geological time period abbreviations are as follows: C: Cretaceous, Pa: 1268 Paleocene, E: Eocene, O: Oligocene, M: Miocene, Pl: Pliocene, Pe: Pleistocene, H: Holocene. 1269 Figure 4. Proposed geographical location of the Black Mountain Corridor (BMC). Inset a) satellite 1270 imagery of the BMC – dashed line denotes the putative location of the barrier; b) elevation (masl); c) 1271 IGBP land cover; d) temperature (driest quarter mean); f) precipitation (driest quarter mean). 1272 1273 Figure 5. Proposed geographical locations of the Burdekin Gap (BG) and Saint Lawrence Gap (StLG). 1274 Inset a) satellite imagery of the BG and StLG – large and small dashed lines denote the putative 1275 locations of the BG and StLG, respectively; b) elevation (masl); c) IGBP land cover; d) temperature 1276 (warmest quarter mean); e) precipitation (driest quarter mean). 1277 1278 Figure 6. Proposed geographical location of the Brisbane Valley Barrier (BVB). Inset a) satellite 1279 imagery of the BVB: the small dashed line denotes the extent of the Brisbane Valley; the Main (1), 1280 McPherson (2) and Border (3) Ranges are indicated by curly brackets. b) IGBP land cover; c) elevation 1281 (masl); d) temperature (warmest quarter mean); e) precipitation (driest quarter mean) overlaid with 1282 substrate moisture index (SMI, annual mean, 75% opacity). 1283 1284 Figure 7. Geographical location of the Hunter Valley Barrier (HVB). Inset a) satellite imagery of the 1285 BVB: extents of the Hunter and Goulburn River Valleys are shown within the bounds of the small and 1286 large dashed lines, respectively; the proposed location of the Cassilis Gap is shown by solid lines. b) 1287 elevation (masl); c) IGBP land cover; d) temperature (wettest and warmest quarter means overlaid, 1288 75% opacity); e) precipitation (driest quarter mean) overlaid with substrate moisture index (SMI, 1289 annual mean, 75% opacity). 1290 1291 Figure 8. Proposed geographical locations of the Southern Transition Zone (STZ) and East Gippsland 1292 Basin (EGB) biogeographical barriers. Inset a) satellite imagery of the STZ and EGB: the area 1293 encompassed by the solid line corresponds to the Illawarra district, the blue dashed line represents 1294 the putative bounds of the Lake George Gate, solid star indicates location of the Tallaganda region; 1295 b) elevation (masl); c) IGBP land cover; d) temperature (driest, wettest, and coldest quarter means 1296 overlaid, 75% opacity); e) precipitation (driest and wettest quarter means overlaid, 75% opacity). 1297